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GROUSE NEWS

Newsletter of the Grouse Group of the

IUCN/SSC-WPA Galliformes Specialist Group

Galliformes Specialist Group

Issue 42 November 2011

Grouse News 42 Newsletter of the Grouse Group

2

Contents From the Editor 3

From the Chair 3

News from the Galliformes Specialist Group 4

Conservation News

U. S. Fish and Wildlife Service lists Cantabrian Capercaillie and five other foreign avian species

under the Endangered Species Act.

5

Research Reports

Research on Russian grouse 7

Landscape connectivity modeling for sharp-tailed grouse and greater sage-grouse – An overview of

current research in Washington State, USA

14

Notes on the Cantabrian capercaillie: winter roosting trees, historical distribution and lack of

knowledge

17

Toxic Scat: A mechanism to prevent overdosing on plant chemicals by grouse 24

Capercaillie lek within the city limit of St. Petersburg, Russia 29

Grouse - Habitat Relationships: monitoring, scale, and management - A short summary of the PhD

Thesis

32

Genetic differentiation of the Western capercaillie highlights the importance of South-eastern

Europe for understanding the species phylogeography

33

Capercaillie (Tetrao urogallus L.), an endangered species due to disturbance and predation, can be

named “the Rose of The Little Prince” in the southern edge of its distribution in Europe

36

Spatial use of capercaillie in the southern Vosges Mountains inferred from non-invasive genetic

sampling

40

Landscape genetics and behavioral ecology of greater prairie-chickens (Tympanunchus cupido) 41

Recent grouse literature 42

Snippets

Pyrenean Network for the mountain Pyrenean galliforms - GALLIPYR Project 50

Desperately seeking stable, 50-year old landscapes with patches and long, wide corridors 51

12th

International Grouse Symposium, Matsumoto, Japan, 20-24 July 2012 52

Jack Connelly receives Hamerstrom Award for his grouse work 52

BLM Developing New Sage-Grouse Conservation Strategy 53

Gateway West Transmission Line Project a Potential Threat to Sage Grouse 54

Article on the effect of wind farms on Cantabrian capercaillie 54

New update of The IUCN Red List 55


3

From the Editor

The earthquake and the following tsunami with the subsequent nuclear crisis stopped the conference in

Japan this summer and forced the organizers to postpone it. As you all remember, it was decided to

postpone it till July 2012. According to Hiroshi Nakamura, the organizer in Japan, it is planned to have

the 12th

International Grouse Symposium in July next year. The Japanese have control over the problems

they had with the nuclear plants and other problems they had. The web site with more information will be

up and go later this autumn.

There are many interesting articles in this issue of Grouse News. An article summarizing many

years of research on grouse in Russia and Norway is published. Contribution dealing with different

aspects of capercaillie is presented like winter roosting trees of Cantabrian capercaillie, genetic

differentiation in western capercaillie, problems with disturbance and predation. It is also articles on

sharp-tailed grouse, greater sage-grouse, and greater prairie-chicken. Research on how grouse prevent

overdosing of plant chemicals in presented. Under snippets sage-grouse conservation strategy and

potential threat, and information on a galliform project is found. We would also extend our

congratulations to Jack Connelly who received the Hamerstrom Award for his grouse work.

In the last issue we had some discussion and early results of using GPS technology for studying

grouse. We had hoped for a follow-up of this topic in this issue from those of you using the GPS

technology, but unfortunately no contributions related to this has come. If you are working with this topic

we ask you to write something for the next issue,

The 6th European conference on black grouse will be held 17-20 September 2012 in Gysinge in

Sweden. An updated web page is soon coming.

Tor Kristian Spidsö, Editor Grouse News

Skilsøtoppen 33, N-4818 Færvik, Norway, [email protected]

Don Wolfe, Co-editor North America

G. M. Sutton Avian Research Center, University of Oklahoma, P.O. Box 2007, Bartlesville, OK 74005,

USA, [email protected]

From the Chair

The next year, 2012, will be a tough one for our international conference travel agendas, with the

International Wildlife Management Congress in Durban, South Africa (9-12 July 2012), the International

Grouse Symposium in Matsumoto, Japan (20-24 July 2012, http://cert.shinshu-

u.ac.jp/eco_lab/modules/tinyD4/), and the 6th

European Conference on Black Grouse in Sweden 17-20

September 2012, http://www.ebc.uu.se/blackgrouse/).

Having both the IGS and the ECBG in the same year might not be ideal, and is a result of

postponing the IGS due to the tsunami and nuclear catastrophe in Japan in 2011. Yet, based on previous

lists of participants, there is normally not much overlap between the IGS and its international attendance

of mostly scientists, and the conservation practice-focused ECBG. I therefore hope for both meetings

being well attended.

Thanks to the dedication of organizer and host professor Hiroshi Nakamura and his team, the

12th

International Grouse Symposium in Matsumoto, Japan, will take place largely as planned, with four

days of conference followed by extended field trips either to the Japanese Alps to experience the

famously tame Japanese rock ptarmigan, or to Hokkaido and the habitats of the Japanese hazel grouse.

The (improved!) IGS website will go online soon to start the registration process. Please help spreading

the information, and particularly also encourage student grousers to attend. I will be there, and I hope to

see many of you in Matsumoto for a stimulating symposium.

Ilse Storch, Chair, Grouse Group within the IUCN-SSC/WPA Galliformes SG (GSG),

Co-Chair, IUCN-SSC/WPA Galliformes SG.

Department of Wildlife Ecology and Management, Institute of Forest Zoology, University of Freiburg, D-

79085 Freiburg, Germany, [email protected]

mailto:[email protected]


http://cert.shinshu-u.ac.jp/eco_lab/modules/tinyD4/

http://cert.shinshu-u.ac.jp/eco_lab/modules/tinyD4/

http://www.ebc.uu.se/blackgrouse/


4

NEWS FROM GALLIFORMES SG

Issue 5 of G@llinformed (Edited by Gilbert Ludwig, Finland) went out in early July 2011. It included a

further technical comment on the utility of satellite tags for tracking grouse. There is also a report on a

WPA field training workshop in Pakistan in April, which was supported by James Goodhart through the

GSG.

At the instigation of the SSC Chair’s Office, we are continuing to discuss how Taxon Advisory

Groups (TAGs) of the European Association of Zoos and Aquaria (EAZA) and IUCN/SSC Specialist

Groups (SGs) could work together more closely for mutual benefit. Longstanding connections with the

EAZA Galliformes-TAG and the World Pheasant Association’s (WPA) European Conservation Breeding

Group (ECBG) pre-adapt the Galliformes SG to forge strong links with zoos, private breeders, along with

our many in situ experts. SSC has now asked us to become a pilot SG for this approach, one goal of

which is to seek EAZA institutional support for the GSG, which would undoubtedly enable us to operate

more efficiently and pro-actively. The other inevitable benefit would be increased interaction between in

situ and ex situ species experts, resulting in better strategic planning of conservation action.

Richard Fuller (Red List focal point, Univ. of Queensland) has contributed for the Galliformes

SG to a current consultation within SSC on the minimum standards of documentation required for

assessment of a species to be valid. The object is to simplify the process.

BirdLife International, as the Red List Authority for all birds, has highlighted a few Galliformes

species for re-assessment of their current threat category: Taiwan partridge, Grey-striped partridge,

Venezuelan wood-quail, Siamese fireback and Sula megapode, the last two being reviewed last year also.

This process is moderated by WPA via the Threatened Galliformes Forum on the BirdLife website. All

GSG members have until 5 December to comment, and to suggest any other species for consideration in

the light of the latest information on status and threats. Among the grouse species, the Siberian spruce

grouse might deserve re-assessment. It is currently listed as near-threatened, and its habitat is rapidly

deteriorating due to commercial forestry. Please get in contact if you should have information on

population and/or habitat that would allow a re-assessment.

GSG Co-Chairs Peter Garson and Ilse Storch and WPA Director Phil McGowan will have a

good opportunity to discuss the changing roles of and relationship between GSG and WPA at the SG

Chairs meeting being organised by IUCN and SSC in Abu Dhabi (23-27 February 2012). We need to

rationalise partners and members and strategically share the workload on behalf of all our 73 threatened

species. The shape of the GSG for 2013-16 (the next IUCN quadrennium) needs to emerge from these

and later discussions.

Peter Garson & Ilse Storch, Co-Chairs, IUCN-SSC/WPA Galliformes SG, October 2011


5

CONSERVATION NEWS

U. S. Fish and Wildlife Service lists Cantabrian Capercaillie and five

other foreign avian species under the Endangered Species Act. Don Wolfe

Recently, Ilse Storch asked me if I would be willing to write a note about this listing action for inclusion

in the Grouse News. Without giving it much thought, I initially agreed, but have since realized that this

listing action is quite a delicate and controversial issue. Rather than expressing strictly my own opinions

and conclusions (likely erroneous), I sent a query out to several grouse biologists in the United States,

some of whom presently work for the U. S. Fish and Wildlife Service, for their comments. Below are a

few of the comments (names have been omitted to protect the guilty). I hope that the comments may

provide some insight, or at least some entertainment. Following the comments received are the press

release by the USFWS on this action, and the official USFWS policy on listing foreign species, which

may help to clear things up.

Comment 1. It isn't April first, is it? I was floored when I read this. I don't know the answer to any of

the questions asked except that this has "Trouble" written all over it.

Comment 2. I would assume that the action was requested by Spain or the IUCN as a measure to prevent

the smuggling of birds that are hunted illegally in Spain. I doubt this would have happened

unless USFWS border agents caught someone trying to do it. This action probably has no

bearing on the listing status of domestic species. . . . they are "apples and oranges". For what it's

worth.

Comment 3. It doesn't make sense that it would be easy to list a Capercaillie and impossible to list a

Gunnison Sage-Grouse.

Comment 4. I'm as puzzled by all of this as (name omitted) is. Perhaps someone in USFWS can shed

some light on these decisions. The grouse conservation issues in this country are befuddled by

bureaucracies. Consider that the new BLM sage-grouse conservation strategy (not that new, just

recycled, it was originally trotted out in 2004) will result in the formation of 17 new teams or

groups or committees and that many of the most knowledgeable people on the subject are

apparently not involved at all. The USFWS seems to be fine with this approach. I am reminded

of a quote from Adlai Stevenson..."There is nothing more horrifying than stupidity in action."

Comment 5. I'm ccing (name omitted) in our DC international listing division, as she will be able to

provide a more detailed response. However, the short answer to the justification for foreign

species is that the ESA, section 4, applies to both foreign and domestic species and Section 8

provides for international cooperation.

USFWS Press Release – 11 August 2011

The U.S. Fish and Wildlife Service (Service) announced today a final rule to protect six foreign bird

species found on islands in French Polynesia and in Europe, Southeast Asia, and Africa as endangered

under the Endangered Species Act (ESA).

The rule implements federal protections provided by the ESA for the Cantabrian capercaillie,

Marquesan imperial pigeon, Eiao Marquesas reed-warbler, greater adjutant, Jerdon’s courser, and slender-

billed curlew. Populations of each of these species are small, fragmented, and declining, making them

particularly vulnerable to genetic threats associated with small populations and extinction.

This determination follows a thorough review of the best available scientific information,

comments from the general public, peer reviews, and any new information received during the public

comment period following publication of the proposed rule to list these species.

Significant threats to these six foreign bird species include habitat loss, overutilization and

inadequate existing regulatory mechanisms. Information on climate change was available for only one

species, the slender-billed curlew; based on this information the Service found climate change to be a

potential threat to this species.

Granting foreign species protection under the ESA means that the import or export of any of the

species, or their parts or products, as well as their sale in interstate or foreign commerce, is prohibited.

Permits for these prohibited actions may be issued for specific purposes consistent with the ESA.

The final rule will publish in the Federal Register on August 11, 2011, and become effective on

September 12, 2011.

http://us.vocuspr.com/Newsroom/Query.aspx?SiteName=fws&Entity=PRAsset&SF_PRAsset_PRAssetID_EQ=128028&XSL=PressRelease&Cache=True


6

Official USFWS policy on listing foreign species

(http://www.fws.gov/endangered/what-we-do/international-activities.html)

The Branch of Foreign Species (BFS), like the domestic listing side of the Endangered Species Program,

determines the status of foreign species and whether they should be added to the Federal list of threatened

and endangered wildlife and plants and receive protections provided by the Endangered Species Act of

1973, as amended. The BFS also assesses foreign species already listed as threatened or endangered to

determine whether they should be reclassified from threatened to endangered, reclassified from

endangered to threatened, or removed from the list (delisted).

Why List Foreign Species?

The Endangered Species Act (ESA) requires the Service to list species as endangered or threatened

regardless of which country the species lives in. Benefits to the species include prohibitions on certain

activities including import, export, take, commercial activity, interstate commerce, and foreign

commerce. By regulating activities, the United States ensures that people under the jurisdiction of the

United States do not contribute to the further decline of listed species. Although the ESA’s prohibitions

regarding listed species apply only to people subject to the jurisdiction of the United States, the ESA can

generate conservation benefits such as increased awareness of listed species, research efforts to address

conservation needs, or funding for in-situ conservation of the species in its range countries. The ESA also

provides for limited financial assistance to develop and manage programs to conserve listed species in

foreign countries, encourages conservation programs for such species, and allows for assistance for

programs, such as personnel and training.

Don Wolfe, G. M. Sutton Avian Research Center, University of Oklahoma, P.O. Box 2007, Bartlesville,

OK 74005, USA [email protected]

http://www.fws.gov/endangered/what-we-do/international-activities.html

http://www.fws.gov/endangered/what-we-do/international-stories.html#prohibition



7

RESEARCH REPORTS

Research on Russian grouse Olav Hjeljord, Per Wegge & Andrey V. Sivkov

A big conservation problem in Europe is dwindling populations of forest grouse. In continental

Europe grouse has disappeared from large areas. Northern populations in Scandinavia and Finland

still inhabit most of their former ranges, but numbers are below what they used to be, and

pronounced cycles and peak populations rarely occur. Causes may be natural or man-made. If the

latter is the problem – direct (habitat destruction) or indirect (increased predation) - the grouse

situation should be different in undisturbed forests. With this as a background we set out to study

grouse in the huge forest reserves of Northern Russia.

Throughout the Russian taiga there is a network of nature reserves (zapovedniks). In Eurasia they

represent the last remaining areas of large, undisturbed, boreal forest, some cover several 1000 km2.

These reserves are research laboratories, were only scientists and reserve staff have legal access.

Extensive monitoring of animal abundance, plant phenology, berry and seed production and climate are

done on a yearly basis. In some reserves such data have been collected for more than 50 years.

During the last 20 years, most intensively during the last 10 years, we have studied capercaillie

(Tetrao urogallus), black grouse (Tetrao tetrix) and hazel grouse (Bonasa bonasia), in Russian forest

reserves. A main objective has been to understand grouse habitat preference and range use in forests

undisturbed by man. Recently, more attention has been on population dynamics. We have worked in the

Pechora Ilich-, the Central Forest- and the Pinezhskiy State reserves. During the latter part of the study,

we selected the Pinezhskiy reserve (421 km2), in the Pinega region, 300 km south-east of Archangelsk, as

most suitable for our work, and most of the data presented here are collected in this reserve. The

Pinezhskiy State reserve has a gentle topography with patches of rugged karst relief. Mature forest

dominate with spruce (51 %) on areas of poor and medium drainage, pure pine stands on elevated ridges

(2%) and mixed forests (11 %). Secondary, younger forest is mostly fire generated (27 %). Part of the

reserve is dominated by large bogs (10 %).

The project has been a collaboration between Norwegian and Russian scientists, and data on

Russian grouse have been compared with similar data collected in Western Europe. During the study 85

capercaillie, 104 black grouse and 40 hazel grouse have been radio collared. Fourteen Norwegian and

Russian students have completed their diploma and Master thesis work on the project.

Cycles still intact in Pinega

There is no straight forward answer to whether there are more or less grouse in the forests of the

Pinezhskiy reserve compared to Scandinavian forests. In preferred habitats, densities of capercaillie and

black grouse are higher but the difference to Scandinavia is not dramatic. The hazel grouse is another

matter, their numbers may be huge (Figure 1). At population peaks hazel grouse may be the most

common bird in the forest, more numerous than thrushes and smaller birds. Historically, the Pinega

region was a centre for export of hazel grouse; during the winter hundreds of sledge loads of the bird went

to the markets in the towns to the south. In St. Petersburg alone the yearly import of hazel grouse is

reported to have been more than 2 million birds in good years. Also capercaillie and black grouse are

common birds of the Russian taiga, and because most grouse-work in Europe has been on these species,

we initially selected the two species as most suitable for comparative studies. Recently we have turned

more of our attention to hazel grouse.


8

Figure 1. Fluctuations in grouse numbers in Pinezhskiy State Reserve, Pinega. Data collection and

population estimates were done by reserve personnel from a modified Kings method.

Grouse habitat use and social organization in Pinega – similarities and differences to Europe

In general, the capercaillie of the Pinezhskiy reserve prefers continuous, older forest while the densest

populations of black grouse are found in areas with large bogs and open landscape (Rolstad et al. 2009).

Habitat preference of hazel grouse has not been analyzed. As we have the most complete data on habitat

use of capercaillie the following discussion will be on this species. We have monitored 42 males and 13

females radio collared on leks.

Use of range. Capercaillie in Northern Russia use their range much like capercaillie in Europe. Birds

occur in subpopulations around the leks (Figure 2). Most males stay within 2-3 km from the lek centre

year around (Lande 2002). Local configuration of preferred summer habitat affects both direction and

movements of birds from the leks in spring (Beshkarev et al. 1995, Hjeljord et al. 2000). Both sexes show

great site fidelity and return to the same habitats in consecutive years. Their mean annual home range is

generally 600-700 ha and the whole lek population of males may use 30-40 km2 over the year. Hens move

more freely in the landscape than males, and some may disperse to nesting areas beyond neighbouring

leks (Lande 2002). They seem

to use the same general areas

both for nesting and brood

movements in succeeding

years (Gregersen & Olstad

2002, Wegge et al. 2005).

Figure 2. Annual range use

by 44 radiotagged cocks on 3

adjacent leks based on 881

cock locations in Pinezhskiy

State Reserve, Pinega. The

lines represent 90 % MCP for

each lek population. Centre of

leks are indicated by a circle

(Lande 2002).


9

Use of habitat. Choice of habitat resembles that of Western European capercaillie. When leaving the lek

in late spring/early summer, male capercaillie search for shaded and humid sites dominated by rich

vegetation of herbs and, interestingly, also by the moss Polytrichum spp (Bjordal & Hillmann 2002). Also

Semenov-Tjan-Sanskij (1960) reported Polytrichum spp. to be part of capercaillie summer diet. A

preference for shaded sites may be due to delayed phenology with higher quality of preferred forage

plants.

Also young broods prefer moist forest types, particularly spruce–swamps and herb–spruce forest

rich in insects and the drier, bilberry-rich Vaccinium spruce forest extensively (Wegge et al 2005).

Caterpillars, probably the most important food for young broods (Picozzi et al. 1999, Wegge & Kastdalen

2008) were twice as abundant in Vaccinium spruce forests than in the other habitat types (Wegge et al.

2005). In the generally moist and insect-rich forest of the Pinezhskiy reserve we would expect rather short

daily movements of broods searching for food. However, average daily movements of broods during the

first 4 weeks were >500 m, similar to that of young broods at Varaldskogen, Norway (Wegge et al. 1982).

This may indicate that predator avoidance rather than search for preferred forage is the main reason for

extensive brood movements (Wegge et al. 2005). In summer, the average distance between male

capercaillie positions when located during two consecutive days was 396 m, also indicating extensive

movements by adults (Bjordal & Hillmann 2002). Partly overlapping home ranges of ca 200 ha were

similar to that reported from Fennoscandia (Rolstad et al. 1988).

Social organization. Most leks are big. Between 20 and 30 males attended the biggest leks in the

Pinezhskiy reserve. In Western Europe there are few leks of this size today. Apparently the large

continuous blocks of old growth forest with little natural disturbance in Russian forest reserves provide

capercaillie with enough time and space to build up large lek populations. In the fragmented and rapidly

changing forests of Western Europe neither time nor habitat is sufficient for the establishment of such

leks. As most copulations on a lek is performed by the highest ranking male, fewer but larger leks may

increase competition for breeding within the population (Rolstad et al. 2009).

On the large leks in the Pinezhskiy reserve daytime home ranges of males were of the same size

(ca 50 ha) and distributed within the same distance from the lek centre (within ca 1 km) as reported

elsewhere (Wegge et al. 2003). However, contrary to what has been found elsewhere (but see Storch

1997) home ranges of the attending birds were almost completely overlapping (Figure 3), but with males

mutually avoiding each other. The clumped spatial distribution may be an adaptation to avoid long

commuting distance to centers of big leks, or a behavioural strategy by subdominants for gaining an

opportunity to mate. The big leks drew birds from rather large areas causing an inter lek distance of

nearly 4 km, compared to around 2-2.5 km reported from Fennoscandia. Probably, during population

peaks, mutual avoidance will push males beyond 1 km from lek centre and contribute to space out leks

maximally, and they remained spaced out also during subsequent population lows (Wegge et al. 2003).

Figure 3. Distribution of daytime ranges of 11 radio-tagged males at a lek with > 25 attending males

during spring 2000 in Pinezhskiy State Reserve, Pinega (Wegge et al. 2003).


10

Logging the northern taiga – the effect on grouse

Comparing grouse populations within the Pinezhskiy reserve with populations in a logged area adjacent

to the reserve, we tested the hypothesis that logging causes an increase in density of generalist predators

and a decrease in reproductive performance of grouse. Grouse were counted along survey lines during

August 1999 and 2001 (Borchtchevski et al. 2003). The somewhat surprising result, however, was a

positive rather than negative effect of logging on grouse reproduction (Figure 4) and no effect on

abundance of small and medium-sized predators, except for red fox, whose tracks were recorded only in

the logged area (Table 1). The estimated total density (adults plus chicks) of capercaillie was generally

lower in the logged area compared to the reserve (3.6 vs 6.2 birds per km2) whereas the opposite was

found for black grouse (7.0 vs 5.4 birds per km2) with no difference in density of hazel grouse (44.3 vs

46.0 birds per km2) (Borchtchevski et al 2003). Tolerance of hazel grouse to logging was also

demonstrated in Komi, southeast of

the Archangelsk region, where

hazel grouse density was similar or

changed only slightly in logged

compared to un-logged areas

(Beshkarev et al. 1995, Romanov

1963), whereas capercaillie

decreased almost three-fold in

abundance in logged forest

(Romanov 1979).

Figure 4. Reproductive success of

grouse inside the Pinezhskiy State

Reserve and in the logged area

outside the reserve, during August

1999 and 2001. Bars show

standard error (Borchtchevski et

al. 2003).

Table 1. Track crossings per 10 km of survey lines per 24 hoursa of carnivorous mammals, mountain hare

and red squirrel. Recordings were made in the Pinezhskiy State Reserve and in the logged area outside

reserve during February 2001 (Borchtchevski et. al. 2003) and in Varaldskogen during January-

February 2006-2010 (J. Rolstad og P. Wegge unpubl. data). Significant differences (p, 0.05) in bold

letters.

Inside Reserve Outside Reserve Varaldskogen Species Tracks/10 km SE Tracks/10 km SE Tracks/10 km SE

Stoat/weasel 4.6 1.91 3.0 0.85 1.3 0.68 Pine marten 0.7b 0.27 0.6 0.39 1.0 0.31 Red fox 0 - 1.0 0.34 3.1 0.94 Wolverine 0 - 0.03 0.03 0 - Otter 0.04 0.04 0 0 0 - Lynx 0.1 0.08 0.3 0.12 0 - Mountain hare 24.6 3.78 53.5 10.03 - - Red squirrel 42.9 7.05 4.3c 0.73 2.8c 1.13 a Badger (Meles meles) not sampled, relatively common in Varaldskogen and absent in Pinega, mountain

hare not sampled in Varaldskogen. b Long time average within the reserve is between 1 and 1.5 tracks/10 km/24 hrs.

c Different from inside reserve.


11

Goshawk as the important predator. A total of 105 remains of dead grouse were found during the survey

(17 per 100 km, the same frequency in logged area and in reserve). From visual inspection, 72 % was

killed by birds of prey, mainly goshawk, the rest by predatory mammals. The proportion killed by the

goshawk was almost identical in the reserve and the logged area, 71 % and 73 %, respectively

(Borchtchevski et al. 2003). These figures compare well with what was found for radio collared birds: 88

% killed by birds of prey (mainly goshawk), 10 % by pine marten and 2 % unknown. At Varaldskogen in

Norway predatory mammals were much more important: 55 % was killed by goshawk, 40 % by red

fox/pine marten and 5 % by unknown predators (Wegge & Rolstad 2011).

Few red foxes in the Russian taiga. Tracks of red fox were recorded only in the logged area. Here we

encountered one fox track/10 km/24 hours (Table 1), compared to 4.4- and 12.3 tracks/10km/24 hours in

northern and southern Finland, respectively

(Kurki et al. 1998), and 3.1/10 km/24 hours in

Varaldskogen, Norway (Table 1) (Wegge &

Rolstad 2011). In the forests of northern Russia,

with loose and deep snow, the red fox is rare

and is mostly found around villages and on

agricultural land (Vaisfeld 1985). Within the

reserve, the red fox has not been recorded

during the last 10 years of track surveys (Rykov

2000). More pine marten tracks were recorded

in Pinega (0.6- and 0.7 tracks/10 km/24 hours in

logged area and in the reserve, respectively)

(Table 1), than in northern Finland (0.3

tracks/10 km/24 hours). However, the long term

average of track counts within the reserve is

between 1.0- and 1.5 tracks/10 km/24 hours,

which is comparable to southern Finland (1.4

tracks/10 km/24 hours) and Varaldskogen (1.0

tracks/10 km/24 hours) (Table1).

Figure 5. The red fox is rare or absent in the

Northern Russian taiga. Either big paws or

broad, moose-skinned skis are required for

travelling on the soft and deep snow.

A “grousefriendly” way of logging? The generally better reproduction of grouse in the logged area

indicates that there are no numerical or functional effects from generalist predators on grouse following

logging and fragmentation. On the contrary, our data indicate higher losses of eggs and chicks to

predation within the old forest of the reserve.

We believe the particular cutting methods applied outside the reserve should be kept in mind

when evaluating these results (Fig. 6). Although logging has caused a coarse-grained forest mosaic

(average patch size varying from 0.5- to 2.8 km2), single trees as well as groups and patches of trees of

little economical value are left within clear cuts and stands of young forest, giving the area a

heterogeneous composition. Approximate age distribution of forest stands is 41 % mature forest (> 80

years), 42 % secondary forest (20-60 years) and 14 % clearcuts and young forest (< 20 years). A factor

contributing to the higher reproductive success of grouse in the logged area may be the populations of

hares being double that recorded in the reserve (Table 1), and possibly acting as a buffer and reducing the

predation on grouse eggs and chicks. Of special interest is a recent study from the same area by

Borchtchevski and Sivkov (2009) indicating that clutch sizes of capercaillie may be larger in the logged

forest compared to the reserve.


12

Figure 6. Selective cutting outside the Pinezhskiy State Reserve, Pinega.

Reproduction and survival of Pinega grouse

The long series of data on yearly change in numbers of grouse, predatory mammals and rodents as well as

climate and fruit/seed production provides a unique opportunity for disentangling the factors causing the

rise and fall of grouse populations in the Pinezhskiy reserve. However, additional data are needed for a

thorough understanding, particularly on the dynamics of reproduction and survival of the Pinega grouse

during the increase and downfall of the populations. We hope to obtain such information by radio

collaring hazel grouse, the species with the widest fluctuation in numbers, and which is presently in the

increase phase. Below we present some preliminary data on grouse population dynamics from the 229

birds we have already radio collared, and from data collected by reserve scientists over many years.

Adult survival appears to be on the same level as in Fennoscandia but may vary considerably

from year to year. Loss of eggs varies between 10- and 50 %, which appears to be lower than in the west.

A comparative study of egg loss in artificial nests indicated a significantly higher loss to predatory

mammals in the logged forest at Varaldskogen, Norway compared to Pinega (Pollen & Ingul 2011,

Wegge et al. subm. ms.). August brood size is usually between 4 and 5 chicks, and does not vary with the

rise and fall of the population, but there is a close correlation (R = 88, p < 0.001) between the proportion

of broodless hens and population change. Data presented in Fig. 1 indicate that recent population peaks of

hazel grouse tend to be on a higher level than before. This coincides with the disappearance of small

rodent cycles and concurrent decrease in populations of stoat and weasel over the past 20 years.

Dramatic population crashes. The most intriguing aspect of the population curve of the Pinega grouse

populations is the sudden crashes of the hazel grouse. While populations build up for 2 or 3 years, the

downfall occurs during a single year. Apparently the crash is caused both by high adult mortality during

the winter following the autumn peak as well as poor reproduction the following summer. During spring

of 2006 (crash year) we followed 8 radio collared females of black grouse intensively. Seven of these

apparently lost their eggs during egglaying or early incubation and were never recorded with chicks. A

pertinent question is whether we see a predator/prey oscillation in the Pinezhskiy reserve: a low rate of

predation permit a high production of chicks in the increase phase, in Western Europe generalist

predators, particularly the red fox, prevents such increases.


13

More data is needed, and a closer investigation is planned on the hazel grouse that we have now

radio collared. Actually the fluctuation of grouse numbers in Pinega resembles those in Norway 50-100

years ago. Several reports from that time indicate sudden crashes when both adult survival and

reproduction failed (Hjeljord 1980).

Acknowledgements

This work was supported by grants from the Norwegian Research Council and the Directorate for Nature

management. We thank Vladimir Borchechevski for fruitful collaboration and John Gunnar Dokk for

valuable assistance in the field.

References

Beshkarev, A. B., Blagovidov, A., Teplov, V. & Hjeljord, O. 1995. Spatial distribution and habitat

preference of male capercaillie in the Pechora-Illych nature Reserve in 1991-92. - In: Jenkins, D.

(ed). Proceedings International Grouse Symposium 6: World Phesant Association, Reading, UK

and Instituto Nazionale per la Fauna Selvatica. Ozzano dell`Emilia, Italy.

Bjordal, H. & Hillmann, C. 2002. Male capercaillie Tetrao urogallus in summer – use of a pristine forest

reserve in northwest Russia. - M. Sc. Thesis, Agricultural University of Norway, Ås. 42 pp.

Borchtchevski, V. & Sivkov, A. 2009. Is the capercaillie (Tetrao urogallus) clutch size density

dependent? - Ornitologia 6: 119-131 (in Russian).

Borchtchevski, V. G., Hjeljord, O., Wegge, P. & Sivkov, A. V. 2003. Does fragmentation by logging

reduce grouse reproductive success in boreal forests? - Wildlife Biology 9: 275-282.

Gregersen, H. & Olstad, T. 2002. Habitat use and movement of capercaillie Tetrao urogallus broods in a

forest reserve in NW Russia during summer. - M. Sc. Thesis, Agricultural University of Norway,

Ås. 44 pp.

Hjeljord, O. 1980. Viltbiologi. Landbruksforlaget. 318 pp.

Hjeljord, O., Wegge, P., Rolstad, J., Ivanova, M. & Beshkarev, B. 2000. Spring-summer movements of

male capercaillie Tetrao urogallus: A test of the ”landscape mosaic” hypothesis. - Wildlife

Biology 6: 251-256.

Kurki, S., Nikula, A., Helle, P. & Lindèn, H. 1998. Abundance of red fox and pine marten in relation to

the composition of boreal forest landscapes. - Journal Animal Ecology 67: 874-886.

Lande, K. 2002. Spacing patterns and annual range use of capercaillie Tetrao urogallus in a pristine taiga

forest in NW Russia. - M. Sc. Thesis, Agricultural University of Ås. 44 pp.

Picozzi, N., Moss, R. & Kortland, K. 1999. Diet and survival of capercaillie (Tetrao urogallus) chicks in

Scotland. - Wildlife Biology 5: 11-23.

Pollen, Oppegaard, V. & Ingul, H. 2011. Using artificial nests as a measure of predator abundance and

predator composition: A comparative study between two contrasting forest areas. - M. Sc. Thesis,

Norwegian University of Life Sciences, Ås. 46 pp.

Rolstad, J., Wegge, P. & Larsen, B.B. 1988. Spacing pattern and habitat use of capercaillie during

summer. - Canadian Journal Zoology 66: 670-679.

Rolstad, J., Wegge, P., Sivkov, A. V., Hjeljord, O. & Storaunet, K. O. 2009. Size and spacing of grouse

leks: comparing capercaillie (Tetrao urogallus) and black grouse (Tetrao tetrix) in two

contrasting Eurasian boreal forest landscapes. - Canadian Journal Zoology 87: 1032-1043.

Romanov, A. N. 1963. Ocherk ekologii ryabchika v Komi ASSR. - Trudi Komi filial Akademii Nauk

SSSR, Siktivkar 12: 7986.

Romanov, A. N. 1979. Obiknovenniy glukhar. - Edition nauka, Moscow. 143 pp.

Rykov, A. M. 2000. Okhotnichi jivotnii. In: Puchina, L. V., Goryachkin, S. V., Glazov, M. V., Rykov, A.

M. & Rykova, S. Y. (Eds); Struktura I dinamika prirodnih komponentov Pinejskogo zapovednika

(Severnaya taiga ETR, Archangelskaya oblast). - Bioraznoobrazie i georaznoobrazie v karstovih

oblastyah, Archangelsk: 179-193.

Semenov-Tjan-Sanskij, O. J. 1960. Die Økologie der Birkhuhnevogel (Tetraoniden). - Trudy

laplandskogo Gosudarstwennogo Zapovednika 5. (Translation no. 106, Swedish Nature Research

Council).

Storch, I. 1997. Male territoriality, female range use, and spatial organization of capercaillie Tetrao

urogallus leks. - Wildlife Biology 5: 187-192.

Vaisfeld, M. A. 1985. Lisica. - In: Nasimovitch, A. A. & Isakov , J. A. (Eds); Pesec, lisica, enotovidnaya

sobaka. Edition Nauka, Moscow: 83-115.

Wegge, P. & Kastdalen, L. 2008. Habitat and diet of young grouse broods: resource partitioning between

capercaillie (Tetrao urogallus) and black grouse (Tetrao terix) in boreal forests. - Journal

Ornithology 149: 237-244.


14

Wegge, P., Storaas, T., Larsen, B.B., Bø, T. & Kolstad, M. 1982. Woodland grouse and moderen forestry

in Norway. A short presentation of a new telemetry project and some preliminary results on

brood movements and habitat preference of capercaillie and black grouse. - Proceedings

International Grouse Symposium 2: 117-122.

Wegge, P., Kvålsgard, T., Hjeljord, O. & Sivkov, A. V. 2003. Spring spacing behaviour of capercailllie

Tetrao urogallus males does not limit numbers at leks. - Wildlife Biology 9: 283-289.

Wegge, P., Olstad, T., Gregersen, H., Hjeljord, O. & Sivkov, A. V. 2005. Capercaillie broods in pristine

boreal forest in northwestern Russia: the importance of insects and cover in habitat selection. -

Canadian Journal Zoology 83: 1547-1555.

Wegge, P. & Rolstad, J. 2011. Clearcutting forestry and Eurasian boreal forest grouse: long-term

monitoring of sympatric capercaillie Tetrao urogallus and black grouse T. tetrix reveals

unexpected effects on their population performances. - Forest Ecology Management 261: 1520-

1529.

Wegge, P., Ingul, H., Pollen, Oppegaard, V., Halvorsrud, E., Hjeljord, O. & Sivkov, A. V. (submitted

ms.). Comparing predation on forest grouse nests by avian and mammalian predators in two

contrasting boreal forest landscapes by use of artificial nests.

Olav Hjeljord, Department of Ecology and Natural Resource Management, Norwegian University of Life

Sciences, Box 5003, 1432 Ås, Norway, [email protected]

Per Wegge, Department of Ecology and Natural Resource Management, Norwegian University of Life

Sciences, Box 5003, 1432 Ås, Norway, [email protected]

Andrey V. Sivkov, State Natural Reserve Pinezhskiy, 123-a Pervomayskaya, Archangelsk Region, 164610

Russia, <[email protected]> <[email protected]>

Landscape connectivity modeling for sharp-tailed grouse and greater

sage-grouse – An overview of current research in Washington State,

USA Leslie Robb and Michael Schroeder Human altered landscapes have impacted the extent to which many grouse populations are fragmented

and isolated, potentially resulting in restricted gene flow and loss of genetic diversity. In Washington

State, sharp-tailed grouse (Tympanuchus phasianellus) and greater sage-grouse (Centrocercus

urophasianus) currently occupy approximately 3% and 8% of their historical ranges, respectively. These

areas are fragmented by habitat loss and tend to be isolated from each other and from occupied habitat in

adjacent states and provinces. Consequently, maintaining and restoring landscape connectivity has been

identified as a key conservation strategy for persistence of these grouse species in the state.

The Washington Wildlife Habitat Connectivity Working Group (WHCWG) is a public-private

partnership of state and federal agencies, universities, tribes, and non-governmental organizations that

was formed in part to address the need for a science based effort to help guide coordinated action to

maintain or improve landscape connectivity for wildlife in Washington State. As part of this effort the

WHCWG recently completed a modeling analysis of landscape connectivity for 16 focal species

(WHCWG, 2010); two of the focal species examined were sharp-tailed grouse and greater sage-grouse.

The connectivity analysis used a cost-weighted distance modeling approach to identify linkages

between areas of occupation (for grouse only). Cost-weighted distance modeling is a novel approach to

understanding potential movement by a species as it moves throughout a landscape that has impediments

and/or barriers created by natural and human related disturbance. In summary, the modeling process uses

spatial data layers to create maps illustrating the ease and extent of movement outward from areas of

interest. For sharp-tailed and greater sage-grouse, these areas were occupied range determined with the

aid of lek surveys, radio telemetry, and habitat evaluation. The spatial data layers used included: 1) land

cover/land use, 2) elevation, 3) slope, 4) housing density, 5) roads, and 6) forest structure. Parameters

associated with each spatial data layer were assigned a resistance score reflecting the difficulty that

particular feature posed to movement relative to “ideal” habitat. For example, for the two grouse species

the parameter “shrub-dominated habitat” had the lowest resistance to movement score while

“urban/developed” the highest. The data layers were then combined and the resistance scores summed for

each grid cell to create a map illustrating the resistance to movement in the landscape for sharp-tailed

grouse and greater sage-grouse respectively. Linkages illustrating the “easiest” movement pathway, in

terms of landscape resistance, between specific pairs of occupied areas (Habitat Concentration Areas in

the figures) were then modeled for both species (Figures 1 and 2).






15

Figure 1. Sharp-tailed grouse linkages in Washington State, USA (WHCWG, 2010).


16

Figure 2. Greater sage-grouse linkages in Washington State, USA (WHCWG, 2010).

The WHCWG has begun a finer scale connectivity analysis of the Columbia Plateau Ecoregion

in eastern Washington and sharp-tailed grouse and greater sage-grouse are included as focal species in

this on-going effort. The Columbia Plateau Ecoregion analysis uses additional data layers and a smaller

grid cell than the statewide analysis (30 m vs. 100 m). This finer scale of resolution will be used to

examine occupied range with respect to landscape resistance as well as potential movement pathways

within occupied areas by modeling linkages between leks. Additionally, the WHCWG is using telemetry,

lek persistence, and genetic data collected from greater sage-grouse in Washington to help validate these


17

connectivity models.

Cost-weighted connectivity modeling products are useful tools for providing insight and

guidance for management and conservation of grouse species. In Washington it is anticipated that these

products will help inform: 1) implementation of agricultural programs to benefit sharp-tailed grouse and

greater sage-grouse, 2) placement of structures such as wind farms and powerlines, 3) identification of

potential reintroduction sites, 4) conservation efforts through the modeling of various restoration

scenarios, and 5) maintenance and enhancement of linkage corridors. The full analysis by the WHCWG

and all mapping products are accessible at http://www.waconnected.org.

References

WASHINGTON WILDLIFE HABITAT CONNECTIVITY WORKING GROUP (WHCWG) 2010.

Washington connected landscapes project: Statewide analysis. - Washington Departments of Fish

and Wildlife, and Transportation, Olympia, Washington, USA. 209 p.

Leslie Robb, P.O. Box 1077, Bridgeport, Washington, 98813, USA, [email protected]

Michael Schroeder, Washington Department of Fish and Wildlife, P.O. Box 1077, Bridgeport, Washington,

USA, [email protected]

Notes on the Cantabrian capercaillie: winter roosting trees, historical

distribution and lack of knowledge Manuel A. González & Vicente Ena

Summary

In winter the capercaillie spend much time roosting to avoid energetic expenditures. This grouse mainly

selects trees that facilitate food, detection of predators and escape flights. Although roosting sites are

broadly documented in other European populations, few data exist on characteristics of Cantabrian

capercaillie roosting sites. We report the first observations of roosting and foraging trees for Cantabrian

capercaillie in a Mediterranean environment. The study area was within the historical distribution of the

Cantabrian capercaillie although it has not been currently considered as occupied habitat by this

subspecies. Scots pines and Pyrenean oaks were used differently by the Cantabrian Capercaillie. We

recommend further research in this Mediterranean habitat for the capercaillie to assess the likely different

habitat use (coniferous and deciduous) and its derived impact on the Cantabrian capercaillie behaviour.

Historical Mediterranean distribution of this grouse should be further considered within any action plan

and ecological studies of this primarily boreal bird.

KEYWORDS: Cantabrian capercaillie, historical distribution, Mediterranean forests, roosting trees,

winter.

Introduction

Among the basic requirements of a bird, the roost site selection is a crucial behaviour with important

fitness bearings (Cody 1981). In temperate latitudes, winter strongly affects survival in non-migrant birds

(Lima et al. 2005), and poses strong challenges to the energy balance and homeostasis of individuals

roosting at night (Walsberg 1986, Wegge et al. 1989). Capercaillie Tetrao urogallus is a big forest grouse

that mainly occurs in boreal and temperate coniferous forests from east Siberia to the Iberian Peninsula

(del Hoyo et al. 1994). It roosts at daylight and night using trees but also snow burrows (Thiel et al.

2007). In winter the main food resource throughout most of its range are pine needles, a superabundant

but low-energy food resource but in the Cantabrian caducifolius forests capercaillie mainly eat understory

plants as ferns, bilberry Vaccinium myrtillus or the evergreen holly Ilex aquifolium and alpine juniper

Juniperus communis subsp. alpina if present (Gjerde and Wegge 1989, Storch 1993, Rodríguez and

Obeso 2000, Blanco-Fontao et al. 2010). To overcome the energetic problem in winter capercaillie spends

much time roosting and it selects needles with a high energy and a low resin content that is decomposed

and digested in the caeca with the aid of bacteria (Lindén et al. 1984, Moss 1983, Spidso and Korsmo

1994). Overall, capercaillie in winter selects trees for both feeding and nocturnal roosting (Spidso and

Korsmo 1994, Storch 1993, Thiel et al. 2007). These wintering sites usually show structural heterogeneity

in tree age, species and size that facilitate food acquisition, detection of predators and escape flights

(Spidso and Korsmo 1994, Storch 1993, Thiel et al. 2007). In Norway most spring roosting sites used by

cocks were located underneath the lower branches in old coniferous trees and in stands with a higher

density of trees than in the surroundings (Finne et al. 2000). In case of high snow-cover and low predator

http://www.waconnected.org/




18

densities, Capercaillie also use snow burrows (Thiel et al. 2007), but this behaviour is rare in capercaillie

populations from central and southern Europe (Marjakangas et al. 1984).

The endangered Cantabrian subspecies Tetrao urogallus cantabricus lives almost entirely in

deciduous forests of the Cantabrian Mountains. In winter it feeds almost exclusively on buds from

deciduous trees as well as leaves of the evergreen holly tree Ilex aquifolium (Obeso 2003, Rodríguez and

Obeso 2000). However, if Scots pine Pinus sylvestris plantations are available then pine needles occur in

the winter diet of the Cantabrian Capercaillie (Rodríguez and Obeso 2000). The main habitats of the

subespecies are Eurosiberian forests dominated by beech Fagus sylvatica, birch Betula pubescens or

sessile oak Quercus petraea (Obeso and Bañuelos 2003), but it also occurs in Pyrenean oak Quercus

pyrenaica forests both in the Eurosiberian (i.e., northern slope of the range) and in the Mediterranean

biogeographic region (i.e., southern slope of the range). This Mediterranean environment is the only

Cantabrian area where pine plantations constitute a relevant percentage (30%) of the forested surface as

well as considered to be the warmest (i.e., two dry months in summer) and the southernmost limit

distribution of capercaillie (González et al. 2010). Up to date 12 leks have been registered in the historical

Cantabrian Capercaillie distribution within the Mediterranean region, however, more leks are likely to be

located with further survey efforts (González unpublished data).

Pyrenean oak is a marcescent oak species more tolerant to xeric conditions than other oaks and

cope better with fire so it can be seen recruiting in slopes covered by Erica australis / aragonensis (i.e.,

those slopes subject to recurrent man-made fires (Luis-Calbuig et al. 2010). In the Cantabrian range,

winter roosting sites are likely to be trees rather than snow burrows since only once an anecdotic snow

burrow was registered in a beech Fagus sylvatica forest when the snow cover exceeded half a meter

permitting snow-roosting (Figure 1; Martínez 1993). Otherwise, observations in Eurosiberian forests (i.e.,

well known to be occupied by the capercaillie) suggest holly as the most

likely roosting and/or foraging tree in winter since it is the only

evergreen tree in these deciduous forests (Blanco-Fontao et al. 2010,

Castroviejo 1975, Rodríguez and Obeso 2000).

The low numbers, secretive behaviour and rough habitat of the

Cantabrian capercaillie makes much difficult to locate its winter roosting

sites. Even more, in winter the leafless trees are not expected to act as

feeding trees for the Cantabrian capercaillie except holly. Here, we

present the first observations on winter roosting and foraging trees of

Cantabrian capercaillie that curiously have been detected in its

Mediterranean distribution.

Figure 1. The only snow burrow documented in the Cantabrian

Mountains, in a Eurosiberian beech Fagus sylvatica forest, snow cover

~1m, year 1982. Photo: Vicente Ena

Methods

Our study occurs in two sites included in an area at the southern slope of the Cantabrian Mountains which

corresponds to the southernmost edge of capercaillie distribution (González et al. 2010). The area is

located in the Mediterranean region bordering to the north with the Eurosiberian region (González et al.

2010). Average temperature ranges between 4ºC and 9ºC and the annual precipitation ranges from 866 to

1,100 mm. Precipitation is unevenly distributed throughout the year, with rain mainly in spring and

autumn and a severe drought for two months during summer, characteristic of the Mediterranean climate

(Blondel and Aronson 1999). The landscape is slightly mountainous (elevation ranges from 800 to 1700

m a.s.l.). Dominant forests are supra-Mediterranean unburned (more than 50 years old) and post-fire

Pyrenean oak forests, which account for the 20% of the area and monospecific Scots pine plantations

younger than 50 years old (8%) (Costa Tenorio et al. 2005). Bilberry is completely absent or very scarce

(<0.5% of the forest ground cover; González et al. 2010).

Remaining 72% of the area is covered by semi-natural habitats mainly composed of Populus

nigra, Fraxinus excelsior and Alnus glutinosa riparian lowland forests, meadows, heaths Erica australis,

brooms Genista sp. and man-made infrastructures such as crops, open mines, roads and villages. Human

density is very low (0.6 people/km²) and population has declined by 85% since the 1950’s (INE 2008).

This decline has lead to the abandonment of pastures with the consequent increase of the forested

landscape (Morán-Ordóñez et al. 2011). The main economic activity is livestock rearing, followed by coal

mining industry, agriculture, forestry and hunting (Penas 1995).


19

The two studied sites are within the 17th

century Cantabrian capercaillie distribution although in

recent literature it has not been considered as occupied by the subspecies (Figure 2; González et al. 2010).

At that time (i.e., 17th

century) the main distribution of the Cantabrian capercaillie was in the

Mediterranean region within the Pyrenean oak stronghold widespread from south of the Cantabrian

mountains to Portugal such as today happens (Madoz 1848, Castroviejo 1975, Martínez 1993). Both

studied sites are within the only area where the Cantabrian capercaillie commonly dwell both in

deciduous and coniferous trees (i.e. Mediterranean distribution; González et al. 2010).

Figure 2. Dark grey historical distribution of Cantabrian capercaillie; light grey current Cantabrian

capercaillie distribution; black dot study area.

From 2005 to 2011 we visited two wintering sites where previously Cantabrian Capercaillie

night dropping groups were frequently detected (González pers. obs.). Distance between both sites was

>10 km, thus we assumed that the birds visiting each winter site were mostly different. One of the sites

consisted of 12 mature (>50 years old) planted Scots pines higher than 15 m occupying 40 m2 surrounded

by a 500 m2 grassland (hereafter pine site; A in Figure 3). South ~20 m of the pine site a small Pyrenean

oak fragment (20 ha) occurs while north ~200 m there is a big oak forest fragment (400 ha). The other site

was a half hectare Pyrenean oak forest fragment (hereafter oak site; B in Figure 3). It was mainly

composed by large oaks higher than 12m in open forest stand with clean undercanopy, surrounded by

secondary cover after repeated fires (e.g. Erica australis heathlands).


20

Figure 3. Forested patches of the study area: dark grey patches known to be occupied by the

Capercaillie; light grey patches not known to be occupied by the Capercaillie; A location of the winter

pines site; B location of the winter oaks site.

From 2005 to 2011 we visited two wintering sites where previously Cantabrian Capercaillie

night dropping groups were frequently detected (González pers. obs.). Distance between both sites was

>10 km, thus we assumed that the birds visiting each winter site were mostly different. One of the sites

consisted of 12 mature (>50 years old) planted Scots pines higher than 15 m occupying 40 m2 surrounded

by a 500 m2 grassland (hereafter pine site; A in Figure 3). South ~20 m of the pine site a small Pyrenean

oak fragment (20 ha) occurs while north ~200 m there is a big oak forest fragment (400 ha). The other site

was a half hectare Pyrenean oak forest fragment (hereafter oak site; B in Figure 3). It was mainly

composed by large oaks higher than 12m in open forest stand with clean undercanopy, surrounded by

secondary cover after repeated fires (e.g. Erica australis heathlands).

Visits consisted of observations of the sites one hour before sunset until dark since it is the time

when the grouse fly to the night sites (Johnsgard 1983; Watson and Moss 2008) and were repeated once a

month from the first snowfall to the end of winter (35 visits to each site). Not to disturb birds, we aimed

to detect capercaillie with a 85 mm 20-60x telescope and 10x40 binoculars from an elevated and

reasonably distant fixed point (>20 m far from the edge site). The day after a visit with zero observations,

an experienced observer completely surveyed the site to homogenize detectability in two different sized

sites, as described in González et al. (2010) in order to find capercaillie night dropping groups. We

considered a positive observation if we observed at least one capercaillie or one group of droppings the

day after.

Figure 4. Capercaillie hen roosting in a Scot

pine in the Cantabrian Mountains. Photo:

Jorge Falagán


21

Results

Each field survey showed a positive observation in both pines (n=35) and oak site (n=35; Table 1).

Capercaillie were always detected landing in the treetops of the pines site and never in the oaks site. After

each negative observation (i.e., oaks site) during the next day at least one droppings group (range 1-12)

was always registered. They were collected not to consider them in the next survey. It showed that the

oaks site was constantly used at least once a month for roosting in winter. Otherwise, in the pines site two

hens were detected in nine occasions, one hen and one cock in two occasions and one cock and two hens

once. In the remaining 17 visits only one solitary hen occurred. Feeding records are incomplete since pine

branches frequently hindered us from adequately observation of the Capercaillie behaviour. However, in

nine occasions we observed birds pecking needles showing these pines act as both foraging and roosting

trees (Table 1).

Table 1. Observation days and birds observed; feeding means any bird seen feeding or not on the pines.

Pines site Oaks site

Date hens (n ) cocks (n ) total (n ) feeding droppings groups (n ) droppings groups (n )

Nov´05 2 0 2 ? 2

Dec´05 2 0 2 Yes 3

Jan´06 2 1 3 ? 3

Feb´06 1 1 2 Yes 5

Mar´06 0 0 0 5 6

Oct´06 2 0 2 Yes 4

Nov´06 2 0 2 No 2

Dec´06 2 0 2 ? 2

Jan´07 2 0 2 ? 8

Feb´07 2 0 2 Yes 12

Mar´07 0 0 0 1 3

Oct´07 2 0 2 ? 6

Nov´07 1 0 1 Yes 9

Dec´07 1 0 1 Yes 11

Jan´08 1 0 1 ? 1

Feb´08 1 0 1 ? 7

Mar´08 0 0 0 5 9

Oct´08 1 0 1 ? 4

Nov´08 2 0 2 ? 6

Dec´08 1 0 1 No 7

Jan´09 1 0 1 ? 3

Feb´09 1 0 1 ? 10

Mar´09 0 0 0 No 6

Oct´09 1 0 1 ? 9

Nov´09 1 0 1 ? 8

Dec´09 1 0 1 ? 3

Jan´10 1 0 1 ? 5

Feb´10 1 0 1 Yes 7

Mar´10 0 0 0 7 11

Oct´10 1 0 1 ? 6

Nov´10 1 1 2 Yes 9

Dec´10 1 0 1 Yes 5

Jan´11 1 0 1 ? 4

Feb´11 1 0 1 ? 3

Mar´11 0 0 0 4 5

N of positive observations 35 35


22

Discussion

In the oak site we never observed capercaillie although signs of night presence were found in each survey.

Otherwise, in the winter pines site we frequently observed capercaillie roosting and feeding. We only had

one replicate per forest type which highly differed in size, however, to our knowledge both sites are the

only monitored and known to be consistently used winter after winter in the Cantabrian range. It does not

imply they build a proper study system, although they are anecdotic reports clearly suggest a different use

between oaks (leafless in winter) and pines trees in winter in the Cantabrian range. This is highlighting

the lack of knowledge on Cantabrian capercaillie ecology, especially winter habitat selection.

In March we neither detected birds in the pines site nor night dropping groups in the oaks site

however we returned twice after the first March date. Hence, the abandonment date of both sites

(February-March) coincides. This date corresponds with the end of winter, last snowfalls and the

beginning of the reproductive period. We speculate about the Cantabrian capercaillie in winter

distinctively use oaks and pines however both of them are abandoned on the same date.

Our five years observations in this Mediterranean area seems to agree with results from central

Europe: Capercaillie in winter faithfully use large coniferous trees for feeding and/or roosting while large

deciduous trees may be used just for roosting likely because leafless. These trees frequently occur in

gentle slopes with open forest stands close to forest edges, while small trees in dense forest stands are

avoided by the Capercaillie (Thiel et al. 2007). Outside this study area, pinewoods and plantations are

very restricted within the Cantabrian capercaillie range and thus they are not likely to constitute an

important resource for roosting and feeding for the entire population. We cannot infer advantages for

capercaillie related to roost in pines or oaks but a different use of them. Otherwise, there is no reason to

benefit pine plantations to improve the winter survival of this grouse which survived without conifers

throughout the Holocene in the Cantabrian mountains (Rodríguez-Muñoz et al. 2010).

Due to the rough habitat of the subspecies, its difficult detectability and low numbers, this is the

first description of two different winter sites for the Cantabrian capercaillie across several years. Although

only two different sample sites were got and our results cannot be extrapolated to the entire Cantabrian

population, this work highlights the different use between them, so further studies should be undertaken

to assess the relative advantages of roosting in pines or deciduous trees in relation to winter-spring

survival and breeding success and so to promote the Cantabrian capercaillie conservation. Taking into

account the Cantabrian capercaillie singularity of living throughout the year in deciduous forests

(Quevedo et al. 2006), the area here described is the only suitable for that assessment. This area is not

only the area of the Cantabrian range where both forest types frequently occur and bilberry, a capercaillie

key species elsewhere (Storch 1995), is near absent (González et al. 2010) but also it may sustain around

40 adult Capercaillie widespread in at least 12 leks (10 % of the total Cantabrian population; own

unpublished data). We recommend considering historical distribution of the Cantabrian capercaillie both

in action plans and ecological studies since requirements of this primarily boreal bird may highly differ in

the underestimated Mediterranean forests of the historical distribution.

Acknowledgements

We are especially grateful to Jose A. López-Garrido, Ramón Balaguer, Fernando Gonzalo, Benito

Fuertes, Emilio de la Calzada, Luis Robles and Jorge Falagán for their invaluable help with the field

observations. Pedro P. Olea comments highly improved this manuscript. BBF was financed by a

predoctoral PFPU-MEC fellowship and MAG by a fellowship of the Universidad de León.

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anti-predator behaviour. - Wildlife Biology 6: 241-249.


23

Gjerde I, Wegge P 1989. Spacing pattern, habitat use and survival of Capercaillie in a fragmented winter

habitat. - Ornis Scandinavica 20: 219-225.

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Manuel A. González & Vicente Ena, Department of Biodiversity and Environmental Management,

University of León, Campus de Vegazana s/n, 24071 León, Spain. [email protected]



24

Toxic Scat: A mechanism to prevent overdosing on plant chemicals by

grouse Jennifer Sorensen Forbey, Graham G. Frye, Xinzhu Pu and John W. Connelly

Introduction

Although abundant and accessible, plants pose significant challenges to herbivores. The high fiber content

and relatively low nutritional value of plants compared to animals makes plants a particularly difficult

food source for birds. For example, birds lack teeth required to reduce particle size and flight limits the

size and complexity of the gastrointestinal tract (Dudley and Vermeij 1992) required for fiber digestion.

These limitations are thought to explain why herbivory is a rare foraging strategy in birds. A less studied

explanation for limited herbivory in birds may also be the diversity of chemical defenses in plants. These

chemical defenses can be toxic to herbivores and energetically costly to detoxify (e.g. Glick and Joslyn

1970, Lindroth and Batzli 1984, Robbins et al. 1987, Guglielmo et al. 1996, Sorensen et al. 2005). Similar

to the limited digestive capacity, flight may limit the size of the liver which is the major organ used to

detoxify and eliminate ingested toxins.

Despite these constraints, most members of the tetraonidae family are herbivores (Sedinger

1997) and many consume plants that are chemically defended. Moreover, many species of grouse are

considered dietary specialists on chemically defended plants that are avoided by other herbivores. For

example, dusky grouse (Dendragapus obscurus) and spruce grouse (Falcipennis canadensis) are winter

specialists on the needles of conifers that contain large concentrations of monoterpenes, a class of toxins

(e.g. Bryant and Kuropat 1980, Boag and Schroeder 1992, Zwickel and Bendell 2005). And ruffed

grouse (Bonasa umbellus) are highly tolerant to the phenolic toxins in aspen (Jakubas et al. 1989, Jakubas

et al. 1993).

How do these avian herbivores

prevent overdosing on potentially toxic

plants? To answer this, we must first

understand the three general fates of

ingested toxins (Figure 1, reviewed in

Sorensen et al. 2006). The first potential

fate of a toxin is to be excreted

unchanged in the feces because it is not

absorbed or metabolized in the gut. The

second potential fate is for toxins to

remain in the gastrointestinal tract where

they can be metabolized by

detoxification enzymes located in

enterocytes of the herbivore or by gut

microflora. Toxins metabolized in the

gut are typically excreted as metabolites

in the feces. The final potential fate is

that toxins are absorbed, delivered to the

systemic circulation, and detoxified in

the liver by detoxification enzymes into

more water-soluble and usually less

toxic metabolites. These water-soluble metabolites are then excreted in the urine. Toxins that enter

circulation and are not metabolized can be delivered to various organs where they may interact with target

sites and cause toxicity. The ultimate goal of the herbivore is to minimize concentrations of toxins in

organs by preventing their absorption and delivery and maximizing their detoxification and excretion.

Although the majority of studies investigating the fate of ingested toxins in herbivores have

focused on mechanisms of detoxification and elimination, we predict that regulated absorption of ingested

toxins is a critical first line of offense for grouse specializing on chemically defended plants. We propose

that mechanisms such as efflux transporters regulate the concentration of toxins that get absorbed into the

bloodstream (Sorensen and Dearing 2006) and reduce reliance on detoxification by the liver. We are

testing this prediction using greater sage-grouse (Centrocercus urophasianus; hereafter sage-grouse).

Sage-grouse provide an extreme example of specialized herbivory in the tetraonids. The sage-grouse is a

dietary specialist on sagebrush (Artemisia spp.). From late fall through early spring, sage-grouse

consume nearly 100% sagebrush, and continue to feed on substantial quantities of sagebrush throughout

the remainder of the year (Rasmussen and Griner 1938, Patterson 1952, Wallestad et al. 1975).

Sagebrush leaves are defended by relatively large quantities of toxins that include monoterpenes,

Figure 1. Simplified diagram of

potential fates of an ingested

plant toxin (represented by X): 1.

Toxins (X) are not absorbed or

metabolized in the gut and are

excreted unchanged (X) in feces.

2. Toxins are not absorbed but

are metabolized in the gut (G) into

new water-soluble metabolites

(XmG) that are excreted in the

feces. 3. Toxins are absorbed and

metabolized in the liver (L) into a

new water-soluble metabolite

(XmL) that is excreted in urine and

then combined with feces in birds.

These potential fates of toxins are

not mutually exclusive


25

sesquiterpene lactones, and phenolic compounds (Shafizad et al. 1974, Welch and Mcarthur 1981, Kelsey

et al. 1982). Many of these compounds are known to have deleterious effects on animals (e.g. Koppel et

al. 1981, Chugh and Jha 2003), yet sage-grouse are totally dependent on sagebrush for forage during

much of the year.

This study is an initial effort to investigate whether regulated absorption is a mechanism of

tolerance to sagebrush chemicals by sage-grouse. We are testing the regulated absorption hypothesis

indirectly by comparing the chemical profile of monoterpenes in the plants consumed at foraging sites to

those excreted in fecal droppings of sage-grouse at these foraging sites. We hypothesized that if sage-

grouse could regulate absorption of ingested monoterpenes from sagebrush, we would see the same

monoterpene profile in plants and excreta.

Methods

We collected browse and fecal samples from sage-grouse in Twin Falls county, Idaho, U.S.A. (4211’ N,

11446’ W). Vegetation in the study area is predominately a mosaic of dwarf sagebrush (black

sagebrush, Artemisia nova; low sagebrush, A. arbuscula) and Wyoming sagebrush (A. tridentata

wyomingensis). We located and flushed radio-collared sage-grouse between 23 January and 14 March

2011. At each flush site, we collected samples of browsed sagebrush plants. Browsed plants were

defined as those of the same species of sagebrush with evidence of at least 10 distinct sage-grouse bite

marks. Clippings from three browsed plants were collected from each flush site and pooled to form a

single browse sample. At each flush site, we collected all fresh sage-grouse fecal droppings and pooled

them to form a single fecal sample. We randomly selected paired fecal and browse samples from six

independent flush sites for this preliminary analysis. We separated leaves from woody stems and debris,

and ground leaf and fecal samples separately in liquid nitrogen prior to analysis.

We used headspace gas chromatography to measure the relative quantity of monoterpenes in the

leaves of sagebrush and excreta of grouse. We sealed 100 mg of wet ground leaf or fecal sample into a 20

ml glass headspace autosampler vial. Samples were incubated at 100° C for 30 min in an Agilent 7694

headspace sampler (Agilent Technology, Santa Clara, CA). One ml of headspace gas was injected into a

J&W DB-5 capillary column (30 m x 250 µm x 0.25 µm) installed in an Agilent 6890N Gas

Chromatogram (GC). Splitless injection mode was used and injector temperature was 250° C. The flow

rate of helium as carrier gas was 1 ml/min. The flame ionization detector was maintained at 300° C.

Chromatographic separation of chemicals was achieved with a temperature program initially held at 40°

C for 2 min and increased to 150° C at 15° C/min and held for 10 min. Monoterpene standards were

prepared by adding 10µl of a mixture of several monoterpenes (50 µg/ml) to a 20 ml glass headspace

autosampler vial and sealed. The standards were analyzed using the same protocol as used for leaf and

fecal samples.

Monoterpene retention times and peak areas for each chemical were calculated by Agilent

ChemStation version B.03.01. For each sample, we selected every major peak that was greater than 1% of

the total area on the chromatogram. We determined the retention times (time detected by the gas

chromatograph detector) and the area under the curve (AUC) for each monoterpene in samples. We

confirmed the identity of monoterpenes in our samples by comparing retention times to those of

monoterpene standards. We quantified the intensity of each peak as AUC for each peak divided by the

dry weight of the sample (g).

We performed a qualitative assessment of chemical similarity by comparing the presence or

absence of chemical peaks from fecal droppings to those from browsed sagebrush within each flush site.

Results

Our preliminary results show that within flush sites, all but the most volatile compounds were found in

both browse and fecal samples (Table 1). Data pooled across foraging sites suggest a relationship

between abundance of monoterpenes in browse samples and abundance of those in fecal samples.

Compounds that were most abundant in plant samples also tended to be most abundant in fecal samples

(Figure 2). Similarly, those that were less abundant in plant samples tended to be less abundant in fecal

samples. However, without knowing the exact amount of leaves sage-grouse consumed and the amount

of feces excreted at a given period of time, it is impossible to determine the percentage of monoterpenes

ingested that were excreted in feces.


26

Retention Time (min)

2.67 4.41 12.64 13.25 13.75 14.36 15.47 17.28 17.46 17.57 18.57 21.61

Foraging Site Unknown 1 Unknown 2 Unknown 3 Unknown 4 Alpha-Pinene Camphene Beta-Pinene Cymene Unknown 5 1,8-Cineole Unknown 6 Camphor

A Plant Plant * * * * * * * * * *

B Plant Plant * * * * * * * * * *

C Plant Plant * * * * * * * * * *

D Plant Plant Feces * * * * * * * * *

E Plant Plant Plant * * * * * * * * *

F Plant Plant Plant * * * * * * * * *

Plant = Compound found only in plant sample

Feces = Compound found only in fecal sample

* = Compound found in both plant and fecal samples

Table 1. Presence of major monoterpenes in plants and fecal droppings from sage-grouse at 6 different

foraging sites (A-F) in south-central Idaho, U.S.A. Retention times (min) refer to the time each

compound (listed as unknown or known monoterpene) was eluted from the gas chromatographer column

and detected. Shorter retention times indicate more volatile compounds

Discussion

Sage-grouse appear to limit systemic exposure to ingested monoterpenes by excreting monoterpenes

unchanged in feces. Toxin excretion is a less explored mechanism that, along with rapid detoxification,

can lower the concentration of toxins in the blood and thereby reduce toxicity (Sorensen et al. 2004,

Sorensen and Dearing 2006). To our knowledge, this is the first study to document that toxin excretion is

a potential mechanism for toxin tolerance in an avian herbivore. In contrast to our results, a previous

study did not detect significant concentrations of monoterpenes in feces collected from the large intestine

of sage-grouse (Welch et al. 1989). However, this earlier study did not investigate fecal droppings from

grouse and the storage of intestinal contents in an intact, slowly frozen and then thawed carcass may have

resulted in volatilization of monoterpenes from the intestine. In addition, the headspace analysis we used

offers a more sensitive assay for detecting and quantifying highly volatile monoterpenes than the analyses

used by Welch et al. (1989).

Figure 2. Mean concentrations (AUC/g dry weight ± SE) of major monoterpenes in browse (plant) and feces from sage-grouse in south-central Idaho. Although substrates differ (plant vs. fecal droppings), similar patterns in relative abundance of monoterpenes are evident across samples types. An exception to the pattern is the most volatile compound detected (Unknown 1), which never occurred in fecal samples.


27

Based on the chemical structure of monoterpenes, these chemicals should be readily absorbed in

the intestine. For example, monoterpenes are highly soluble in blood (Falk et al. 1990) and less than 5%

of either inhaled or orally ingested monoterpenes are typically recovered unchanged in the feces of

mammals and often never detected (Igimi et al. 1974, Foley et al. 1987, McLean et al. 1999, Boyle et al.

2000). Our results suggest that sage-grouse possess mechanisms that actively prevent absorption of

ingested monoterpenes. We propose that monoterpenes are transported out of the gut epithelium and into

the gut lumen by various efflux transporter proteins (Sorensen and Dearing 2006) that are highly

conserved across species (Annilo et al. 2003). Our results are consistent with a study on woodrats

(Neotoma spp.) where ingested alpha-pinene, a monoterpene, was excreted unchanged in feces (Sorensen

et al. 2004). The majority of studies investigating mechanisms of tolerance to toxins in herbivores have

ignored regulated absorption. Our results suggest that this mechanism deserves further attention.

Although there is a strong pattern suggesting that sage-grouse can limit the absorption of many

of the monoterpens in sagebrush, there are several limitations to our study. First, we did not measure the

actual amount of monoterpenes consumed and the amount excreted unchanged and therefore cannot

determine the percentage of ingested monoterpenes that were excreted. Second, additional studies are

needed to identify why the more volatile monoterpenes in sagebrush (with retention times at 2.67 and

4.41 min) did not appear in feces. Rates of volatilization from the upper gastrointestinal tract (White et

al. 1982) and the extent of metabolism in the gut should be investigated. Both of these require feeding

studies with captive grouse that are logistically difficult. Finally, our results are restricted to a single

population of sage-grouse in a single season. The extent of excretion of ingested toxins may be

dependent on season, the chemotype of sagebrush consumed or developmental stage of sage-grouse and

should be further explored.

Understanding the influence of tolerance to toxins by grouse may be important in efforts to

decipher and predict habitat use patterns. For example, grouse at our study site, selectively used dwarf

sagebrush habitats for foraging. The concentrations of toxins in dwarf sagebrush are markedly different

from those in the alternative sagebrush species available on the study area (unpubl. data). Different

chemotypes may influence the extent of absorption and detoxification by sage-grouse and therefore limit

intake of a particular sagebrush taxa. Moreover, climate change may influence tolerance to toxins by

herbivores and further restrict use of certain plant species. For example, rises in CO2 are predicted to

increase concentrations of toxins (Lindroth 2010), and toxins are likely to become more toxic with

increasing temperatures (Gordon 2005, Dearing et al. 2008). Understanding the mechanisms that

facilitate tolerance to toxins may help us predict how changes in land use regimes and climate will affect

the ability of grouse to thrive on a given food source.

The regulated absorption hypothesis offers a novel explanation for the relatively high tolerance

of tetraonids to chemically defended plants. We recommend developing a collaborative effort to

investigate the extent to which regulated absorption plays a role in tetraonid foraging ecology. Such an

effort requires integration of ecological knowledge of diverse plant-tetraonid systems with the

bioanalytical insight and instrumentation available in our lab. The most informative systems will be those

in which birds consume large amounts of potentially toxic plants (e.g. plants generally avoided by other

herbivores). Plants defended by volatile or aromatic toxins (e.g. monoterpenes), such as conifers, are

preferable as headspace analysis can easily be applied to these systems. Monoterpenes represent a

lipophilic class of toxins that would require regulated absorption to be excreted, and comparisons can be

made between these new species and our results from sage-grouse. However, we can measure a range of

other chemical classes of interest (e.g. phenolics, di- and tri-terpenoids, alkaloids, etc). Initial efforts

would require collecting browsed plants and fresh feces from foraging sites and then storing these

samples immediately on ice or at -20C until they can be analyzed for chemical concentration. We

welcome those interested in this type of collaborative effort to contact us.

Acknowledgements

We would thank David Musil, Lisa Cross, and Torrey Ritter for their time and energy trapping, flushing,

locating and helping with all field aspects of this project. Thanks to Richard Lowell and the Magic Valley

Steelhead Hatchery for providing lodging during the field work. We also thank Amy Ulappa for initiating

the sage-grouse research in the lab. Research was supported by the Bureau of Land Management CESU

award No LO9AC15385 and the Idaho Sage-grouse Advisory Committee Office of Species Conservation

Award to J Forbey as well as Boise State University and the Idaho Department of Fish and Game.

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Jennifer Sorensen Forbey, Graham G. Frye and Xinzhu Pu, Department of Biological Sciences, Boise

State University, Boise, ID 83725, USA, [email protected].

John W. Connelly, Idaho Department of Fish and Game, Blackfoot, ID 83221, USA,

[email protected].

Capercaillie lek within the city limit of St. Petersburg, Russia Roald Potapov

I started my bird life studies near the suburban village Komarovo (known in Finnish as Kellomiaki) in

1947, when I was a 14-year-old schoolboy, and I continue to do so to this very day. Komarovo is situated

42 km north from the center of St.-Petersburg, in the southern part of the Karelian Isthmus and now is a

part of the Greater St. Petersburg administrative area. It is surrounded by mixed and coniferous forests

alternating with little bogs and clearings (former Finnish hayfields) being taken over by thickets. The flat

relief alternates with long hills (former glacial moraines) with little bogs and lakes between them. In

1950-1970’s, all these place were populated by a large number of hazel grouse, black grouse, and

capercaillie. A large black grouse lek functioned at a distance of 500-700 meters from the houses at the

village border. With the increase of the human population and of its activity (most importantly both legal

and illegal hunting), the number of these game birds gradually decreased. In 1985, black grouse

disappeared completely from the surroundings of Komarovo. Other grouse (hazel grouse and capercaillie)

decreased in number by several times and it was hardly possible to find them at distances closer than 4-5

km from the village. However, the game birds’ population stabilized owing to the complete prohibition of

hunting in the wide suburban zone around St.-Petersburg, and especially because of the development of a

negative attitude in society to hunters and to hunting in general. In 1950-1970’s it was very common to

meet a hunter with a gun in Komarovo’s streets, however at present such encounters are simply

impossible. As a result, the reverse process of recovery was gradually developing. Hazel grouse were the

first; they appeared again in suitable forest habitats close to Komarovo. In 1979, I found a capercaillie

nest at a distance of about 2 km from the nearest house, and in the following spring, I found a small

capercaillie lek (3 males only) close to this place. During my winter skiing excursions, I began to find

capercaillie snow-burrows, not frequently, but more or less regularly. At last, in January 18, 1993, I found

a place in the pine forest with traces of feeding left by one or two males; they had evidently fed there on

pine needles in pine crowns; I also found 2 fresh female snow burrows located at a distance of 3 m one

from the other. It was only 10 A.M. early morning, and the number of fecal droppings in snow burrows

(74 and 77) with the usual portion of frozen caeca’s excrements demonstrated that the birds spent nearly

14 hours there (that is usual for this time and this ambient temperature constituting -24° C); these females

obviously left their snow burrows undisturbed. The presence of the popular and intensively used skiing

route along a high-voltage powerline 200 meters away from this lek site, as well as its location at a

distance of 550 meters from the nearest houses was very unusual. Later, in March, I visited this place

several times and every time I recognized traces of capercaillie feeding sites in so-called “fodder-pines”.

As it is well known, males prefer to feed on needles of ill or stressed pines around bogs or in marshy

soils, where, because of disturbances in metabolism, such needles offer a higher nutritional value, with a

larger amount of protein and with a smaller amount of calcium (Pulliainen, 1970). Birds will consume up

to 50% of needles from such trees; this intense consumption leads to the appearance of new sprouts and,




30

as a result, tree tops become flattened (Lobachev & Tscherbakov 1936; Potapov 1985). This shape is

retained for a very long time and allows the recognition of capercaillie’s prior activity in places where

these birds have since disappeared. Exactly this shape of pines is characteristic of the mentioned place;

tree crowns were obviously changed by feeding capercaillie. At that time, I thought that this place is only

a place suitable for bird’s feeding; I could not even imagine that it was a real lek site. Because of this, in

the end of March, it was a real surprise for me to see traces of 2 males that crossed the high-voltage line

covered by deep snow and entered the forest in the direction of the place described above. In this place,

traces were finished by clear prints in the snow of clear nuptial activity with walking with partly spread

and lowered wings and flutter-jumps. The followed observations in April – May supported my first

impression: it was a real capercaillie lek.

This place was typical of capercaillie’s lek in this region of the southern taiga belt: it was a

slightly swamped plot of pine forest; this plot was shaped as a long 300 m wide strip between spruce

forests growing in more elevated ground. This plot differed from the neighboring areas of the pine forest

in a sparser arrangement of trees and in the absence of the spruce undergrowth. Therefore, this place

possessed a possibility of circus observation for a distance of 30 – 40 m. The presence of a large number

of tubers, green leaves and winter flower buds of the cotton-grass (Eriophorum vaginatum) was another

important feature. After a prolonged winter diet, this sedge is very important for capercaillie females for

clutch production because of its extremely high nutritional value (Andreev, 1978; E. Potapov, 1993); as

soon as the snow cover disappears prior to the breeding season; females seek for places rich in the cotton-

grass and feed there intensively. In Leningrad District, capercaillie lek sites are associated with places

rich in cotton-grass, presumably because the plant attracts females, and the females attract males to the

lek (Potapov 2008). It is interesting to note one feature of female’s food. After the consumption of such

cotton-grass buds, females produce specific excrements that can be easily found, detecting these places. I

found them constantly. Their excrements look like those of male, but they are sufficiently smaller,

shorter, with a thickened rounded end, and with a white urine patch, without any traces of pine needles;

they also possess a monotonous consistence of a dark green color. All these excrements look fresh at any

time (Figure 1), but, by contrast to male excreta that can stay on the ground for a very long time, female

excrements disappeared rapidly. I performed some experiments: marked such excrements with little sticks

to check up their condition later, but I could not find most of them even the next day (only 20%-30% of

old excreta were found; all the other

were new ones). Those that preserved

till the next day had not changed in

weight, consistency, or color.

However, this observation needs

further studies. The great importance

of the cotton-grass was shown in

tundra ecosystems for populations of

reindeer and especially for lemmings,

including its great role in lemming’s

cycles (E. Potapov, 1993).

Figure 1. Dropping of a female

capercaillie

Among my observations of that period, one was especially exciting. It happened in the evening

of May 1. The snow was virtually gone. A single male visited the lek grounds this evening. The bird

arrived immediately after sunset, landed in the crown on one of pines, uttered the usual raucous “u – au”

(so-called Belching Canto), and was sitting motionless, listening and looking around (above) attentively

during several minutes. After that, it performed the first demonstrative flight with loud wing beats to

another pine at a distance of nearly 30 m. During the following 2 hours, this male performed several such

flights and finished the last of them in the pine right above my shelter where I sat in my sleeping sack.

This male landed on a large thick branch in the lower part of the crown, walked slowly along it and time

and time ate bundles of needles. It began to sing in almost complete darkness, one song followed another.

Intervals between songs sometimes decreased, sometimes increased. Songs became very active after the

moon rise and the black silhouette of the male at the background of the full moon was a unique show. The

uniqueness of this situation was strengthened by music, songs, joyful cries and woman’s laughter that


31

were heard from the nearest cottage where the inhabitants celebrated the Mayday Holiday. Capercaillie

did not pay attention to such unusual, but not so loud sounds, and continued singing until midnight. It

slept soundless and recommenced its singing at 3.30 AM in full darkness. Two females appeared there

about one hour later, in the beginning of the dawn, uttered their usual “ak…ak…ak” and after that the

male stopped its singing and flew down to the ground immediately, to the center of the lek ground, where

it began to sing again, alternating singing with flutter jumps. Judging by sounds, females approached the

male, but subsequent events were invisible for me. Nevertheless, I had no doubts that despite the presence

of only a single male in the lek, it fulfilled its reproductive function effectively.

Regular observations, performed by me since this time till nowadays, demonstrated that

capercaillie regularly used this place for feeding on pine needles and for reproductive courtship activity in

spring. As a rule, only a single old male and 2 – 3 females appeared in this lek. Only in 1998, in the first

days of May, I observed the second male, only a yearling, who walked in a nearly complete lekking

posture with his tail fully open and lifted vertically, but not approaching the old male. Narrow tail feathers

of this young capercaillie in its open and lifted tail looked like spokes in a wheel.

Another problem: how one can explain the existence of the capercaillie lek in a heavy populated

area, in this case in suburbs of such a large megapolis as St. Petersburg? By the way, officially, this place

belongs to the territory of the city. There are two main explanations. First of all, it is the complete

prohibition of any hunting and preservation of forests as around suburban villages, so between marginal

cottages. Fragmentation of forest landscape by local roads and numerous high-voltage lines do not affect

bird habits and behavior seriously. A constant presence of solitary pedestrians gathering berries and

mushrooms in forests during summer seasons is not dangerous for capercaillie; birds avoided disturbance

from such persons easily due to their fine auditory and visual abilities: they simply flew to calmer places.

Problems may arise only with broods, but, as a rule, females place their nests deeper in the forest in

marshy places, only occasionally or never visited by humans; at the beginning of the berry and mushroom

season ( -the end of June), Capercaillie chicks are big enough to avoid disturbance from peaceful visitors.

In any case, this situation provides evidence that capercaillie, the symbol of the wilderness in the remote

past, can dwell very close to humans. It is most unusual when one of the most wild and shy species, such

as capercaillie, move the place of its most private and intimate ritual – the lek – close to heavily build up

area inside the borders of the largest -Russian city Greater St-Petersburg!

There is some other evidence of such possibility. Sergey Kirpichev, one of the best experts of

capercaillie’s life, who is the only person in Russia able to breed successfully this bird in captivity,

informed our colleagues about a wonderful story that had happened in the 1970’s. In a single well-

documented case, in Kamchatka, a new forestry settlement was established deep within a larch forest

during winter. When spring came, the local black-billed capercaillie (T. parvirostris) birds began to

gather, because this place was their traditional lek site. The forestry staff did what they could to minimize

disturbance and the birds quickly became accustomed to them, ceased to be afraid and continued their

activity with courtship displays and singing, occasionally with humans standing within distances of only

3–4 m. Now Sergey Kirpichev still studies capercaillie in a little village (Tver Province north from

Moscow) without any help from Russian Game and Department officials. He believes that capercaillie

possess all the necessary features to be the usual element of the modern heavily changed landscape and its

future depends only on humankind.

References.

Andreev V. N. (ed.) 1978. Seasonal and weather dynamics of phytomass in the subarctic tundra. -

Novosibirsk, 191 pp. (in Russian).

Lobashev S. V., Scherbakov F. A. 1936. The natural forage of Capercaillie during ayear. - Zool. J., 40,

(2): 307-320. (in Russian).

Potapov E., 1993. Ecology and energetics of Rough-legged Buzzard in the Kolyma River Lowlands. -

Unpublished PhD Thesis, Edward Grey Institue for Field Ornithology, Oxford University.

Potapov R.L. 1985. Order Galliformes, family Tetraonidae. - Fauna of the USSR, new series, No. 133.

638 pp. (in Russian).

Potapov R. L. 2008. A lek of the Capercaillie, Tetrao urogallus, near suburban settlement Komarovo on

the Karelian Istmus. - Russian Journal of Ornithology vol.17, Express-issue No. 440: 1400-1406

(in Russian with English resume).

Pulliainen E., 1970. Composition and selection of food in the winter by capercaillie in north-east Finnish

Lapland. - Suomen Riista 22: 67-73.

Roald PotapovZoological Institute, Russian Academy of Sciences, Universitetskaya Enb., 1, St.

Petersburg, Russia. [email protected].



32

Grouse - Habitat Relationships: monitoring, scale, and management -

A short summary of the PhD Thesis Unni Støbet Lande

Capercaillie (Tetrao urogallus) and black grouse (Lyrurus tetrix) have declined over the last 20 – 30 years

in central Europe, United Kingdom and parts of Fennoscandia (Storch 2007, Kurki et al. 2000). Bag

statistics of willow ptarmigan (Lagopus lagopus) show a similar decline in Norway (Statistics Norway

2010). All grouse species are subject to recreational hunting in Fennoscandia and management aims to

ensure sustainable use. To be able to identify factors causing population decline and initiate proper

management actions, a well-functioning monitoring program and management are essential.

In this thesis I have used datasets on capercaillie, black grouse and willow ptarmigan (Fig.1).

The datasets were sampled through large scale monitoring programs by volunteers. The volunteers were

mainly hunters that already had field-experience and knowledge of grouse, and could do the work at low

cost and with few logistical problems. ). In Sweden, transect counts since 1960 exists (Hörnell-

Willebrand 2005). In Norway, bag statistics from each county have been collected since 1972 (Statistics

Norway 2010).

Distance sampling of willow ptarmigan in Norway and Sweden are conducted in slightly

different ways, which makes the inference of the results from wildlife monitoring more challenging.The

results showed that using a GPS to measure perpendicular distances less than 50m from line transects

reduced the reliability of the density estimates. Adding covariates such as habitat (open or closed), and

flushed by dog or handler, improved the models of density estimates. In non-systematic sampling designs

where hunters were free to go

wherever they wanted during

sampling, I found little bias in the

habitat selection of hunters compared

to that of capercaillie and black

grouse, probably caused by the

diversity in habitat use among the

hunters. This indicates that such

sampling of data can also provide

unbiased indices of population

density, given a sufficiently high

number of hunters is used for

monitoring.

Based on the adult density

and chick production from the

density estimates, I found a positive

relationship between the

heterogeneity of the landscape and

both adult density and chick

production of capercaillie and black

grouse. Forest of moderate to high

productivity had a positive effect on

adult density of both species while

contrary to expectation the

proportion of old forest seemed to

have a negative effect on chick

production of both species. This is

supported by findings of a negative

relationship between old forest and

capercaillie in Finland (Miettinen et

al. 2008). Capercaillie and black

grouse are described as synchronous

in time and space (Kvasnes et al.

2010). It seems quite reasonable to

consider an active management for

both species rather than single

Figur 1. The geographic locations of all areas linked to each paper (I-

IV) in this thesis.


33

species management regimes. I did not detect any clear relationship between willow ptarmigan and

habitat at either the individual or the landscape scale. However, at the population scale habitat

relationships were detected.

Together, these results suggest that methods for monitoring grouse species are quite reliable, but

could be improved by rather simple additions. Moreover, habitat composition seems to generate

differences in demographic rates and population densities among populations, but these are not detectable

at the individual or landscape scale. This suggests that management of grouse species should be done at a

scale that captures the heterogeneity in the landscape, most likely over larger areas.

References

Hörnell-Willebrand, M. (2005). Temporal and spatial dynamics of willow grouse Lagopus lagopus.

Doctoral Thesis. Swedish University of Agricultural Sciences.

Kurki, S., Nikula, A., Helle, P., and Lindén, H. (2000). Landscape fragmentation and forest composition

effects on grouse breeding success in boreal forests. Ecology 81: 1985–1997.

Kvasnes, M.A.J., Storaas, T., Pedersen, H.C., Bjørk, S., Nilsen, E.B. 2010. Spatial dynamics of

Norwegian tetraonid populations. Ecol. Res. 25(2):367–374.

Miettinen, J., Helle, P., Nikula, A., and Niemelä, P. (2008). Large-scale landscape composition and

capercaillie (Tetrao urogallus) density in Finland. Ann. Zoo. Fenn. 45:161–173.

Statistics Norway (2010) Jegere på jakt 2009/10 Available at: http://www.ssb.no/jakt_fiske/. Accessed

November 19. 2010 (In Norwegian)

Storch, I. (2007). Grouse Status Survey and Conservation Action Plan 2006-2010. WPA/BirdLife/SSC

Grouse Specialist Group. IUCN, Gland, Switzerland and Cambridge, UK and the World

Pheasant Association, Reading, UK. Online

Lande, U.S. 2011. Grouse – Habitat Relationships: Monitoring, Scale and Management. – Ph.D.

Dissertation, Swedish University of Agricultural Sciences. Opponent: Professor Ilse Storch

Unni Støbet Lande, Norwegian Institute for Agricultural and Environmental Research, Gunnars Veg 6, N

– 6630 Tingvoll, Norway, [email protected]

Genetic differentiation of the Western capercaillie highlights the

importance of South-eastern Europe for understanding the species

phylogeography Marko Bajc, Miran Čas, Tine Grebenc & Hojka Kraigher

Summary Western capercaillie (Tetrao urogallus L.) is a grouse species of open boreal or high altitude forests of

Eurasia. It is endangered throughout most mountain range habitat areas in Europe (Storch 2007).

Considered to be an umbrella species, Western capercaillie serves as an indicator of sufficiently preserved

structures of old coniferous or mixed forests. Genetic differentiation studies of Western capercaillie in

Eurasia (Duriez et al. 2007, Rodríguez-Muñoz et al. 2007, Segelbacher and Piertney 2007, Segelbacher et

al. 2003, Liukkonen-Anttila et al. 2004) revealed the existence of two main genetically identifiable

lineages: the southern and the boreal lineage. The southern lineage has been discussed as a glacial relict

(Duriez et al. 2007, Rodríguez-Muñoz et al. 2007) and has only been described at southern edges of the

species distribution range in Europe. We addressed the question of genetic differentiation of Western

capercaillie populations across all major habitat areas in the Balkans including Rhodope and Rila

Mountains in Bulgaria, Dinaric Mountains (Slovenia, Croatia, Bosnia and Herzegovina, Montenegro,

Serbia) and Southeast Alps in Slovenia. The contact zone between the two lineages and the resulting

implications for conservation strategies in this so far under-studied area of distribution had not been

determined previously.

The findings of an extensive genetic differentiation study based on analysis of mitochondrial

DNA control region one (CRI) sequences of 319 samples revealed the existence and distribution of both

Western capercaillie genetic lineages in the Balkans and Southeast Alps (Figure 1). Slovenian Alpine

population was composed exclusively of boreal lineage, Dinaric population of both, but predominantly

(96%) of boreal lineage and Rhodope-Rila populations predominantly (>90%) of southern lineage

individuals. The Bulgarian mountains were, therefore, identified as the core area of the southern lineage,

and Dinaric Mountains the western contact zone between both lineages in the Balkans. The Balkans

region served as a refugium during the last glacial maximum and is as such of great importance for

understanding phylogeography and genetic differentiation of the species. Genetic lineage composition of



34

Bulgarian and Dinaric capercaillie populations can be interpreted as similar to isolated Cantabrian

(southern lineage) and Pyrenean (contact zone) populations (Duriez et al. 2007, Rodríguez-Muñoz et al.

2007, Segelbacher et al. 2003), respectively. According to high pairwise FST values, the results of

Monmonier's maximum difference algorithm and AMOVA the Bulgarian populations appeared

genetically distinct from Dinaric and Alpine populations. In contrast to Dinaric and Alpine populations

Bulgarian Western capercaillie also exhibited characteristics of a long-term stationary population.

Furthermore, the presence of unique Dinaric southern lineage haplotypes and high levels of genetic

differentiation between the Dinaric and Bulgarian populations suggest that the two populations have been

effectively isolated for considerable time and that the presence of southern lineage individuals in the

Dinarides is very likely not a result of recent long-range dispersal events or illegal release of individuals

from Bulgaria. Our results (Bajc et al. 2011) support the hypothesis that Bulgarian capercaillie should be

considered as a glacial relict and potentially a distinct subspecies (de Juana 1994) and evolutionary

significant unit (Duriez et al. 2007, Rodríguez-Muñoz et al. 2007). Furthermore, phylogenetic analysis

produced evidence of two closest inter-lineage haplotypes, discovered to date, originating from the same

geographic location (Bulgaria), suggesting the possibility that the boreal lineage had evolved from the

southern lineage in the Balkans.

Figure 1. Map of T. urogallus sampling localities in the Balkans and Southeast Alps and haplotype

composition per sampled countries/mountain ranges. Slovenian Alps are shaded blue, Dinarides red and

Rhodope-Rila-Pirin Mountains grey. Black dots mark the localities where only boreal lineage individuals

were discovered, white square markers in black outline represent localities where also southern lineage

individuals were discovered. Each of the 30 unique mitochondrial DNA control region one haplotypes

identified in this study is represented in a different colour: boreal lineage haplotypes in grey tones,

southern lineage haplotypes in red tones. Country and mountain range names and number of sampled

individuals are given for each haplotype composition bar chart. Dashed line marked β represents the only

major genetic barrier detected in the studied region according to Monmonier’s maximum difference

algorithm. Alb: Albania, At: Austria, BiH: Bosnia and Herzegovina, Bg: Bulgaria, Cro: Croatia, Cz:

Czech Republic, D: Germany, Gr: Greece, Hun: Hungary, It: Italy, Kos: Kosovo, Mac: Former Yugoslav

Republic of Macedonia, Md: Moldova, Mne: Montenegro, Rom: Romania, Sk: Slovakia, Slo: Slovenia,

Srb: Serbia, Tk: Turkey, Ukr: Ukraine; sampled countries codes are underlined.


35

The results are presented in the context of conservation of the species in the studied region, its

principal threats and legal protection status. The influences of past forest use and differences in

characteristics of natural habitats of the studied mountain ranges are discussed as important when

interpreting the observed differences in genetic diversities and demographic histories of Western

capercaillie populations. Although all of the studied populations had suffered a decline in the past (Čas

2006, Čas 2010, Adamič et al. 2006, Gačić et al. 2009, Zubić 2009, Petrov 2008), we identified the

Dinaric Western capercaillie as particularly vulnerable to continuing population decline which may

negatively affect the fitness of the population and ultimately its survival as this population exhibited

significantly lower levels of genetic diversity compared to the neighbouring Alpine and Bulgarian

populations. Reduction of genetic diversity has been reported previously for isolated and fragmented

Western capercaillie populations (Segelbacher and Piertney 2007, Segelbacher and Storch 2002). Adding

to the vulnerability of Dinaric Western capercaillie is the fact, that Dinaric population is functionally

isolated due to the loss of connectedness with the Alpine population in Slovenia in 1960s (Adamič 1987,

Čas 2006). The results of our study underline the fact that potential conservation strategies should

consider the existence of the two Western capercaillie lineages and their contact zone, and support the

specificities of the populations in the studied region.

For detailed information see the original paper: Bajc, M., Čas, M., Ballian, D., Kunovac, S.,

Zubić, G., Grubešić, M., Zhelev, P., Paule, L., Grebenc, T. & Kraigher, H. 2011. Genetic differentiation

of the Western capercaillie highlights the importance of South-eastern Europe for understanding the

species phylogeography. - PLoS ONE 6(8): e23602. doi:10.1371/journal.pone.0023602 available at:

http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0023602

Acknowledgments

Foremost we would like to thank all of the co-authors of the original paper published in PLoS One. We

would also like to thank the following colleagues for their invaluable contribution to our research:

Metodija Velevski for providing the information on the status of the Western Capercaillie in Macedonia;

Blažo Jokanović for providing samples from Montenegro; Prof. Dr. Krešimir Krapinec for his selfless

help with collecting samples in Croatia; Dr. Roman Volosyanchuk for additional material from Ukraine

(not included in the analysis); and others who provided samples for analysis: Prof. Dr. Dragan Gačić,

Serbia; Marko Čaleta, Bosnia and Herzegovina; Dr. Stoyan Stoyanov, Bulgaria; Marko Jonozovič, Janko

Mehle, Miha Marenče Jr. and forest rangers from the Slovenian Forest Service; members of the Slovenian

Hunting Association and its professional chairman Srečko Žerjav; Boris Leskovic, the editor of the

Slovenian journal Lovec (The Hunter); Miha Marenče Sr. and park rangers from the Triglav National

Park. We would also like to extend our sincere gratitude to the reviewers and PLoS One staff, whose

constructive comments and suggestions helped improve our paper.

References Adamič, M. 1987. The ecology of capercaillie (Tetrao urogallus L.) in Slovenia. - Strokovna in

znanstvena dela 93. Univerza Edvarda Kardelja v Ljubljani, VDO Biotehniška fakulteta, VTOZD

za gozdarstvo, Ljubljana. Slovenia, 93 p.

Adamič, M., Rapaić, Ž., Popović, Z., Kunovac, S., Koprivica, M., Soldo, V., Marković, B., Maunaga, R.,

Mićević, M. & Ilić, V. 2006. Ugrožene vrste divljači u Bosni i Hercegovini. - Projekat razvoja i

zaštite šuma, Finalni izveštaj, Banja luka, Bosna i Hercegovina, 138 p.

Bajc, M., Čas, M., Ballian, D., Kunovac, S., Zubić, G., Grubešić, M., Zhelev, P., Paule, L., Grebenc, T. &

Kraigher, H. 2011. Genetic differentiation of the Western capercaillie highlights the importance

of South-eastern Europe for understanding the species phylogeography. - PLoS ONE 6(8):

e23602. doi:10.1371/journal.pone.0023602.

Čas, M. 2006. Fluctuation of capercaillie (Tetrao urogallus L.) population in relation to past land use and

forest structures in the South-East Alps. - PhD thesis, University of Ljubljana, Ljubljana,

Slovenia, 263 p.

Čas, M. 2010. Disturbances and predation at capercaillie leks in Alps and Dinaric Mountains. Šumarski

list 9-10: 487–495.

de Juana, E. 1994. Family Tetraonidae. - In: del Hoyo, J., Elliot, A. & Sargatal, J. (eds.). Handbook of the

birds of the World. Lynx Edición, Barcelona, Spain, pp. 376–410.

Duriez, O., Sachet, J.M., Menoni, E., Miquel, C. & Taberlet, P. 2007. Phylogeography of the capercaillie

in Eurasia: what is the conservation status in the Pyrenees and Cantabrian mounts? -

Conservation Genetics 8: 513–526.

Gačić, D., Puzović, S. & Zubić, G. 2009. Capercaillie (Tetrao urogallus) in Serbia – Principal threats and

conservation measures. - Šumarstvo 61: 155–167.

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36

Liukkonen-Anttila, T., Rätti, O., Kvist, L., Helle, P. & Orell, M. 2004. Lack of genetic structuring and

subspecies differentiation in the capercaillie (Tetrao urogallus) in Finland. - Annales Zoologici

Fennici 41: 619-633.

Petrov, I.K. 2008. Distribution and numbers of capercaillie, Tetrao urogallus L., 1785 (Aves:

Galliformes) in Bulgaria. - Acta Zoologica Bulgarica Supplement 2: 35–40.

Rodríguez-Munoz, R., Mirol, P.M., Segelbacher, G., Fernández, A. & Tregenza, T. 2007. Genetic

differentiation of an endangered capercaillie (Tetrao urogallus) population at the Southern edge

of the species range. - Conservation Genetics 8: 659–670.

Segelbacher, G., Hoglund, J. & Storch, I. 2003. From connectivity to isolation: genetic consequences of

population fragmentation in capercaillie across Europe. - Molecular Ecology 12: 1773–1780.

Segelbacher, G. & Piertney, S. 2007. Phylogeography of the European capercaillie (Tetrao urogallus) and

its implications for conservation. - Journal of Ornithology 148: 269–274.

Segelbacher, G. & Storch, I. 2002. Capercaillie in the Alps: genetic evidence of metapopulation structure

and population decline. - Molecular Ecology 11: 1669–1677.

Storch, I. (compiled and edited by) 2007. Grouse: Status survey and conservation action plan 2006–2010.

- IUCN, Gland, Switzerland and World Pheasant Association, Fordingbridge, UK, 114 p.

Zubić, G. 2009. Density and vulnerability of capercaillie (Tetrao urogallus) in the area of Vitoroga in the

Republika Srpska. - Bulletin of the Faculty of Forestry 100: 71–84.

Marko Bajc, Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana, Slovenia, EU.

[email protected].

Miran Čas (Corresponding author), Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana, Slovenia,

EU. [email protected].

Tine Grebenc Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana, Slovenia, EU.

[email protected].

Hojka Kraigher, Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana, Slovenia, EU.

[email protected]

Capercaillie (Tetrao urogallus L.), an endangered species due to

disturbance and predation, can be named “the Rose of The Little

Prince” in the southern edge of its distribution in Europe Miran Čas Introductory reflection

The recognition of causes for endangerment of rare species of wildlife animals, plants or fungi, is the first

step for conservation of a species, including its genetic, ecosystem and landscape diversity and richness.

The recognition of conditions for their optimal habitat and population structures are the second step, and

analysis of each disturbance or habitat-population parameter is the third step for sustainable dynamical

conservation and management of these species in time and space.

Adapting wildlife management for rare species conservation at the edge of their distribution area

need special monitoring and sensitive research approaches between a thin interval of common parameters

in all three steps. The Slovenian Alps and Dinaric mountains are south-eastern edge of the interrupted

Western capercaillie habitats, representing an altitudinal corridor between the fragmented edge habitats in

mixed or coniferous forests of spruce, beech and silver fir through the mountain chains between the

Central and South-eastern Europe (Adamič 1987; Čas and Adamič, 1998; Čas 2001, 2006). The

capercaillie in this area is a rare and endangered species. In Slovenia its hunting is prohibited by the

Slovenian Hunting Association since 1984, and is protected by law since 1993 (Official Gazette of RS,

1993/57).

Distribution of habitats in the Alps is fragmented (Storch 1999, 2007, Čas 2001, 2006), which is

predicted with topography, land use history and climate change (Čas and Adamič, 1998, 2007, Čas 2006,

Kutnar and Kobler 2011), causing alteration of forest habitat (Čas and Adamič 1998, Čas 2002, 2006), as

was confirmed in studies in some similar mountain areas of Europe (e.g. Graf et al. 2007, Braunisch and

Suchant 2008, Anić et al. 2009).

Care and attention for capercaillie conservation in the temperate European mountain ranges

requires great efforts by enthusiasts, environmentalists, biologists, foresters and hunters (e.g. Klaus and

Bergmann 1994, Storch 1999, 2002, Čas 2001, Bollman et al. 2005, Quevedo et al. 2006, Raguž and

Grubešić 2006, Zubić 2009). Just as the knowledge and concern for the conservation of different species

and forest biodiversity differs in different countries around the world, so also differs conservation of






37

capercaillie populations. In some countries or their regions continuous or sporadic monitoring of birds

number at leks and research of capercaillie populations ecology and demography are conducted, (e.g.

Cattadori and Hudson 2000, Čas 2002, Pollo et al. 2005, Storch 2007, Graf et al. 2007, Petrov 2008,

Gačić et al. 2009), as is in some boreal lands (e.g. Rolstad and Wegge 1987, Beškarev et al. 1995, Kurki

et al. 2000, Angelstam 2004). Capercaillie is a protected species in most countries of the region even

though its distribution and population density is not always known, while in some countries it remains

unprotected, and is exposed to hunting, without knowing whether it is at risk or already reaching

extinction (Čas 2010, Bajc et al. 2011). Therefore, the IUCN-GSG with their actions and initiatives are an

important bridging organization between countries, promoting the needs for recording the status and

conservation management strategies of forest grouse populations, and Grouse News is a mean for their

successful deployment.

In some protected areas leks of capercaillie are under permanent supervision of foresters,

environmentalists, biologists and hunters, with appropriate approaches for the species survival which is of

a non-profit mutual interest (Čas 2008). Most of capercaillie leks are monitored by enthusiasts from

different professions, mostly hunters and foresters, who watch over the suitability of the habitat, its

disorders and pressures, and survival and rejuvenation of the sub-populations.

In these mountainous areas of the fragmented habitat boundary of southern Europe the Western

capercaillie occurs as a rare and charismatic species, and each male or female is beloved as the “Rose

of The Little Prince”. With the abundance of large populations of capercaillie in the Boreal forest in the wide area of northern Euro-Asia, where it is an important game species, or in other countries with

different habits and tradition, it may be difficult to understand our attention and precision to review leks

and sub-populations.

Through this reflection we present the article: Disturbances and predation on capercaillie at

leks in Alps and Dinaric mountains (Čas, 2010), which was published in Šumarski list no. 9-10,

CXXXIV (2010), 487-495, as an initiative of members of the GROUSE SPECIALIST GRUP of IUCN in

2008. Publication of research results in this article is important for our orientation in the first step of

studies of its vulnerability and ecology, necessary for the conservation management of habitats and

populations of capercaillie in the Slovenian Alps and the wider area of the Alpine and Dinaric mountains.

Summary Capercaillie (Tetrao urogallus L.) populations in Central and South-eastern Europe cover fragmented

edge habitats and are recorded to be in decline since 1960ies (Klaus and Bergman 1994, Čas 2006).

Survival of sub-populations is sensitive especially in period of winter and spring lek habitats occupancy

and activity (e.g. Rolstad and Wegge 1987, Storch 1999, Elliason and Wegge 2007, Thiel et al. 2007, Čas

2010). Capercaillie leks in Slovenia are present at the south-eastern edge of the Alpine meta-population

and at the north-western edge of Dinarics (Adamič 1987; Čas 2006). These populations were monitored at

leks in two periods in 1980 (466 monitored leks) and 2000 (599). All leks were monitored by local

specialists (hunters and/or foresters) and main causes of observed lek populations decline were addressed

to each endangered lek. Special emphasis was given to predation at leks, as suggested by D. Jenkins

(2008). Among the six identified causes in 1980ies which affected 39 leks the most pronounced were

logging of old-growth forests (at 71.8% of leks) and construction of forest roads (7.7%). In 2000 nine

causes affected 92 leks: (i) mountain tourism (26.1%), (ii) cutting of old-growth forests (19.60%), (iii)

predators attacks (18.5%), (iv) forest management in spring time (9.8%), (v) pastures of livestock with

wire fences in forests (6.5%),(vi and vii) berries picking and overgrowth of abandoned pastures by

forest,(viii) constructions of forest roads and (ix) infrastructure. The most profound change in reasons

between 1980 and 2000 mapping data were: predation at leks, mountain tourism development, and

increase of forest management in spring time, uncontrolled grazing of cattle and sheep in forests,

overgrowth of abandoned pastures into a forest-landscape. A comparison of the increasing percentage of

leks endangered by predators since 1980 has shown positive correlations with increase of the main

predator populations’ densities. Population density of martens (Martes sp.) and wild boar (Sus scrofa)

increased for 150% since 1980, while red fox (Vulpes vulpes) density increased only after 1990. Our

results confirmed the assessment of causes for threats to leks based on descriptions and experiences of

observers as a suitable approach for capercaillie habitat risk assessment. Conclusions from past decline

and negative impacts on lek habitats are important guidelines for foresters and wildlife managers

concerning sustainable forest management practices for maintenance of vital capercaillie populations.

Conclusions and suggestions for adapted forest and wildlife management

A sustainable and dynamic prevalent share of mountain old-growth mixed forests, a moderate road

density and unaggressive and controlled mountain tourism was important for conservation of capercaillie

habitat suitability. Control of the number and influence of predators (by their hunting) (Budiansky 1995,


38

Angelstam et al. 2001, Saniga 2002, Čas 2006, 2010, Sandercock et al. 2010) in the capercaillie lek areas

and the nature of coherent population density of predators were crucial regulators of stable grouse

densities in mountain forest landscapes of Central and South-Eastern Europe (Storch et al. 2005; Čas

2006). This analysis showed that the assessment of the causes of threats to leks on the basis of the

descriptions and experiences of observers are a good indicator of the causes of risks to habitats. Our

results on the current situation and differences regarding the negative impacts on habitats have resulted in

an important guideline for forest and hunting management planning, and for a sustainable multipurpose

landscape use with a continuous presence of forest grouse species.

For detailed information see the original paper: Čas, M. 2010. Disturbances and predation on

capercaillie at leks in Alps and Dinaric mountains. - Šumarski list 9-10: 487-495.

Acknowledgement

The text preparation was supported by the Research Programme Forest Biology, Ecology and Technology

(P4-0107), financed by the Slovenian Research Agency and V4 0175109 (1998-2000), V4-0435 (2001-

2005), V4-0492 (2008-2011) and V4-0497 (2008-2010) research projects funded by the Ministry of

Agriculture, Forestry and Food of the Republic of Slovenia (RS), the Ministry for Science and

Technology of the RS / the Slovenian Research Agency, and the Ministry of the Environment and Spatial

Planning of the RS. We thank all hunters and foresters from the Slovenian Hunting Association (SHA)

with the head of professional services Srečko Žerjav and the Slovenian Forestry Service (SFS) with the

head team of Marko Jonozovič, Janko Mehle and Miha Marenče for their efforts in monitoring the

numbers of capercaillies and leks observations. We would like to thank Prof. Miha Adamič from

Biotechnical faculty, Dep. of forestry in Ljubljana and to Prof. Marijan Grubešić and Prof. Krešimir

Krapinec from Forestry faculty in Zagreb for their support in evaluating research, Dr. Primož Kmecl and

Cilka Zupančič and especially to Dr. Tine Grebenc and Prof. Hojka Kraigher from the Slovenian Forestry

Institute for a fruitful cooperation and support of our research efforts.

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Angelstam, P. 2004. Habitat thresholds and effects of forest landscape change on the distribution and

abundance of black grouse and capercaillie. Ecol Bull 51:173-187

Angelstam, P., M. Breuss, Cas, M., Fischer, I., Ploner, R., Storch, I., R. Summers & A. Zeitler, 2001. Did

rural hunting on generalist predators in past make life easier for grouse? Peca - Črna na

Koroškem, Slovenia. Draft of the potential project (unpublish) : 7 p.

Anić, I., Vukelić, J., Mikac, S., Bakšić, D. & Ugarković, D. 2009. Effects of global climate change on the

ecological niche of silver fir (Abies alba Mill.) in Croatia. Šumarski list 3–4: 135–144.

Bajc, M., Čas, M., Ballian, D., Kunovac, S., Zubić, G., Grubešić, M., Zhelev, P., Paule, L., Grebenc, T. &

Kraigher, H. 2011. Genetic differentiation of the Western capercaillie highlights the importance

of South-eastern Europe for understanding the species phylogeography. - PLoS ONE 6(8):

e23602. doi:10.1371/journal.pone.0023602.

Beškarev, A., Blagovidov. A., Teplov, V. & Hjeljord, O. 1995. Spatial distribution and habitat preference

of male Capercaillie in the Pechora-Illich Nature Reserve in 1991 and 1992. The Sixth

International Grouse Symposium, Udine, p. 48-53.

Bollman, K., Weibel, P. & Graf, R.F. 2005. An analysis of central Alpine capercaillie spring habitat at the

forest stand scale. For Ecol Manage 215:307-318.

Braunisch, V. & Suchant, R. 2008. Using ecological forest site mapping for long-term habitat suitability

assessments in wildlife conservation - Demonstrated for capercaillie (Tetrao urogallus). For Ecol

Manage 256:1209-1221.

Budiansky, S. 1995. Nature’s keeepers : the new science of nature management. New York, Free Press :

310 p.

Cattadori, I. M., Hudson, P. J. 2000. Are grouse populations unstable at the southern end of their range?

Wildlife Biology, (6) 4: 213-218.

Čas, M. 2001. Capercaillie in Slovenia - indicator of devastation, use, development and biodiversity of

mountain forest ecosystems. Gozd. vestn., 59 (10): 411-428 (in Slovene with Engish summary).

Čas, M. 2002. Forest land biodiversity use, degradation and development, co-natural silviculture and

capercaillie (Tetrao urogallus L.) as indicator in Slovenian Alps: research report. Grouse news

24:10-13.


39

Čas , M. 2006. Fluctuation of Capercaillie (Tetrao urogallus L.) population in relation to past land use

and forest structures in the South-East Alps. PhD Thesis, University of Ljubljana (in Slovene

with Engish summary).

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ecosystems, but introduction of a hunting ban in the Slovenian Alps has highlighted certain

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Čas, M. 2010. Disturbances and predation on capercaillie at leks in Alps and Dinaric mountains. -

Šumarski list CXXXIV: 9-10: 487-495.

Čas, M. & Adamič, M. 1998. The influence of forest alteration on the distribution of capercaillie (Tetrao

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Engish summary).

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urogallus L.) population in the Slovene south-east Alps. In Jurc M (ed) Climate changes, impact

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English summary).

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lekking ground in spring. J. Avian Biol. 38: 37–43.

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scenarios in Slovenia. Šumarski list CXXXV, 3–4: 113-126.

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Cantabrian capercaillie Tetrao urogallus cantabricus at leks on the southern slope of the

Cantabrian Mountains, north-west Spain. Bird Cons Int 15, 397-409.

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fragmentation. Oecologia 72 (3): 389–394.

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Galliformes) in Bulgaria. - Acta Zoologica Bulgarica Supplement 2: 35–40.

Raguž, D. & Grubešić, M. 2006. The study of population stability of capercaillie - Tetrao urogallus L. In:

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Quevedo, M., Banue los , M. J., Saez , O. & Obeso, J. R. 2006. Habitat selection by Cantabrian

capercaillie Tetrao urogallus cantabricus at the edge of the species’ distribution. Wildl Biol 12:

267–276.

Sandercock, B.K., Nilsen, E.B., Brøseth, H. & Pedersen, H.C. 2011. Is hunting mortality additive or

compensatory to natural mortality? Effects of experimental harvest on the survival and cause-

specific mortality of willow ptarmigan. J Anim Ecol 80: 244–258.

Saniga, M., 2002. Nest loss and chick mortality in capercaillie (Tetrao urogallus) and hazel grouse

(Bonasa bonasia) in West Carpathians. Folia Zoologica : 202-214.

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Management, RieS Druck und Verlags, München.

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Capercaillie Numbers? Conservation ecology 6, 1 : 6.

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farmland mosaics of central Europe. Landsc Ecol 20: 927–940.

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- IUCN, Gland, Switzerland and World Pheasant Association, Fordingbridge, UK, 114 p.

Thiel , D., Menoni, E., Brenot, J. F. & Jenni , L. 2007. Effects of recreation and hunting on flushing

distance of capercaillie. Journal of Wildlife Management, 71 (6): 1784–1792.

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Republika Srpska. - Bulletin of the Faculty of Forestry 100: 71–84.

Miran Čas, Slovenian Forestry Institute, Večna pot 2, 1000 Ljubljana, Slovenia, EU.

[email protected].



40

Spatial use of capercaillie in the southern Vosges Mountains inferred

from non-invasive genetic sampling. Arnaud Hurstel & Gwenaël Jacob

We report about the first results of capercaillie genetic monitoring in the Vosges Mountains, as a part of

the European LIFE+ project “Des forêts pour le Grand Tétras”. We set the non-invasive monitoring of the

local capercaillie population with the aim to monitor the fragmentation of the species’ habitat and identify

movements of individuals within the Vosges Mountains.

The results presented here are based on the analysis of 256 faeces samples collected from early

February to end of May 2010. Fresh faecal samples were collected in airtight tubes filled with silica gel

and stored frozen upon analysis. After DNA extraction, samples were genotyped at 12 microsatellite loci

(Jacob et al. 2009, Höglund et al., in prep) and an additional locus used to identify the sex of the

individuals (Griffiths, 1998). Each marker was amplified 4–12 times to control for genotyping errors.

Overall, 226 samples (88 %) provided a good-quality genotype that could be used in subsequent analyses

and among those, we identified 71 unique genotypes. The probability of two individuals sharing the same

genotype was low (PIsib = 3.9 x10-3

), which indicates we can confidently distinguish among individuals

based on their genotypes. Thirty-one individuals were identified as females, and 39 as males. The sex of

the remaining individual could not be determined yet. We estimated a number of individuals in the range

76–106 (mean 88) based on genetic capture–recapture methods (program CAPWIRE, Miller et al. 2005).

Forty-four individuals were identified in more than one location. The distance covered within a

day (Euclidean distance between sample locations) ranged from 0.2 to 1.4 km. Summed over the time

period considered in the study, individual movements reached up to 4.2 km in 45 days. These movements

occurred within individual home ranges, during prospection for food or other daily activities. We also

observed movements of nine individuals (13 % of all the individuals) over longer distances, four

individuals moved to areas at a distance of 3.4 –5.2 km, four moved distances of 9.4–14.3 km and one

moved over 27 km.

Given the narrow time window and small number of records, the figures presented above are not

representative of individual home ranges. Being primarily interested in identifying routes used during

movements within the Vosges Mountains, the possibility to detect and track most individuals within the

population makes genetic monitoring a method of choice to achieve this objective. Samples were

collected in 2011 but were not analysed yet. Including a second year of data will provide further

information on spatial use by capercaillie in this area. Hopefully, we will be able to identify preferred

routes and key areas (stepping-stones) used during individual movements. This information will later be

used to implement management measures.

Aknowledgement

We thank all the people who collected samples in the field.

References

Griffiths, R, Double, M.C., Orr, K., Dawson, R.J.G., 1998. A DNA test to sex most birds. - Molecular

Ecology, 7(8): 1071-1075.

Höglund, J., Rudh, A., Thörngren, H. et al. Cross-species amplification of chicken microsatellites and the

development of a multiplex microsatellite protocol for grouse (Tetraoninae). - in prep

Jacob, G., Debrunner, R., Gugerli, F., Schmid, B., Bollmann, K., 2010. Field surveys of capercaillie

(Tetrao urogallus) in the Swiss Alps underestimated local abundance of the species as revealed

by genetic analyses of non-invasive samples. - Conservation Genetics 11(1): 33-44.

Miller, C.R., Joyce, P. & Waits, L.P., 2005. A new method for estimating the size of small populations

from genetic mark−recapture data. - Molecular Ecology 14: 1991-2005.

Arnaud Hurstel, Groupe Tétras Vosges, 1, cour de l’Abbaye, F-68140 Munster, France,

[email protected]

Gwenaël Jacob, University of Fribourg, Unit of Ecology and Evolution, Chemin du Musée 10, CH-1700

Fribourg, Switzerland, [email protected]




41

Landscape genetics and behavioral ecology of greater prairie-chickens

(Tympanunchus cupido) Andrew J. Gregory Ph. D. Dissertation

Kansas State University

Abstract Anthropogenic activities and climate change have dramatically altered landscapes worldwide. The ability

of species to cope and adapt to ongoing changes is likely a function of their behavior, movements, and

sensitivity to fragmentation. Greater Prairie-Chickens (GPC) are a lek mating grouse native to the Great

Plains Landscape Conservation Cooperative (GPLCC), for which inbreeding depression and

anthropogenic avoidance are a concern. The goals of my dissertation were to: 1) identify genetic

correlates of male performance which may influence population viability under current land use practices,

2) identify GPC habitat characteristics and delineate areas of critical GPC habitat necessary for GPC

conservation, and 3) identify the relative importance of distance and habitat quality for maintaining

genetic connectivity among spatially structured populations. First, I found male reproductive success and

survival to be positively associated with genetic diversity. Using multistate modeling in Program Mark,

male survival across the observed range of variation in number of alleles (15-22) increased more than

fourfold from 0.17 to 0.77. Second, I found 35-40% of Kansas, and 1.5 % (11,000 Km2) of the GPLCC,

were considered high-quality lek habitats. Top performing logistic models predicting lek presence

(wi=0.95) included strong effects of grassland cover and avoidance of anthropogenic disturbance. When

this model was applied to putative future landscapes based on climate change and current land use trends

over a 70-year period, I found a 27-40% reduction in habitat area and a 137 Km southeast shift in habitat

distribution. Under equilibrium conditions we expect isolation by distance (IBD) to explain the

distribution of genetic diversity. However, if the landscape restricts dispersal, then we might observe

isolation by resistance (IBR). I used model selection procedures to choose among competing IBR or IBD

models to explain the distribution of genetic diversity among GPC populations across Kansas and the

GPLCC. IBD was never supported (R2<0.02, P>0.09). The best models for Kansas (R2=0.69, P<0.02)

and for the GPLCC (R2=0.46, P<0.02) indicated that human-mediated landscape changes have influenced

landscape permeability for dispersal. The integration of behavioral, landscape, and genetic data provided

new insights on prairie-chicken ecology, and is a powerful approach for developing conservation

strategies for sensitive species.

Location of three samples assigned to a female

capercaillie and collected on the same day

(25.03.2010). All samples were collected within

a 100 m radius.

Approximately 40 days later (05.05.2010), a

fourth sample assigned to the same female is

found 27 km eastward.


42

RECENT GROUSE LITERATURE For a complete bibliography on grouse, go to: http://www.suttoncenter.org/pages/publications

(please note that the link in previous editions may not be current).

Adam, A., L. M. I. Webster, W. Mullen, L. F. Keller, and P. C. D. Johnson. 2011. Quantifying

fenbendazole and its metabolites in self-medicating wild Red Grouse Lagopus lagopus scoticus

using an HPLC–MS–MS approach. Veterinary Parasitology 177:383-386.

Alda, F., P. Sastre, P. J. De La Cruz-Cardiel, and I. Doadrio. 2011. Population genetics of the

endangered Cantabrian Capercaillie in northern Spain. Animal Conservation 14:249-260.

Andersen, O., B. P. Kalterborn, H. C. Pedersen, T. Storaas, H. Solvang, P. F. Moa, and B. R. Hagen.

2010. Undersøkelse blant jaktrettighetshavere i Rypeforvaltningsprosjektet 2006-2011.

Datagrunnlag og noen sentrale funn. [Survey among managers in the Grouse management

project 2006 - 2011. Data and key findings.] NINA Report 433. 32 pp. (in Norwegian with

English abstract). (Willow Ptarmigan).

Augustine, J. K., J. J. Millspaugh, and B. K. Sandercock. 2011. Testosterone mediates mating success in

Greater Prairie-Chickens. Pp. 195-208 in B. K. Sandercock, K. Martin, and G. Segelbacher

(editors). Ecology, conservation, and management of grouse. Studies in Avian Biology (no.

39), University of California Press, Berkeley, CA.

Baines, D., N. Aebischer, M. Brown, and A. MacLeod. 2011. Analysis of Capercaillie brood count data:

long term analysis. Scottish Natural Heritage Commissioned Report 435:1-35.

Baines, D., N. Aebischer, and A. MacLeod. 2011. Analysis of Capercaillie brood count data: habitat

associations. Scottish Natural Heritage Commissioned Report 343:1-14.

Baines, D., N. Aebischer, A. MacLeod, and J. Woods. 2011. Assessing the activity of predators in

relation to Capercaillie hen densities and breeding performance. Scottish Natural Heritage

Commissioned Report 413:1-33.

Bajc, M., M. Cas, D. Ballian, S. Kunovac, G. Zubic, M. Grubesic, P. Zhelev, L. Paule, T. Grebenc, and H.

Kraigher. 2011. Genetic differentiation of the Western Capercaillie – highlights the importance

of South-Eastern Europe for understanding the species phylogeography. PLoS ONE 6(8):

e23602. doi:10.1371/journal.pone.0023602

Baker, W. L. 2011. Chapter eleven – Pre-Euro-American and recent fire in sagebrush ecosystems. Pp.

185-201 IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and

conservation of a landscape species and its habitats. Studies in Avian Biology No. 38.

University of California Press. 646pp.

Barnagaud, J.-Y., P. A. Crochet, Y. Magnani, A. B. Laurent, E. Menoni, C. Novoa, and O. Giminez.

2011. Short-term response to the North Atlantic Oscillation but no long-term effects of climate

change on the reproductive success of an alpine bird. Journal of Ornithology 52: 631-641.

(Black Grouse).

Beck, J. L., J. G. Klein, J. Wright, Justin, and K. P. Wolfley. 2011. Potential and pitfalls of prescribed

burning big sagebrush habitat to enhance nesting and early brood-rearing habitats for Greater

Sage-Grouse. Natural Resources and Environmental Issues 16(5).

http://digitalcommons.usu.edu/nrei/vol16/iss1/5

Beever, E. A., and C. L. Aldridge. 2011. Chapter fourteen – Influences of free-roaming equids on

sagebrush ecosystems, with a focus on Greater Sage-Grouse. Pp. 273-290 IN: S. T. Knick and J.

W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and conservation of a landscape

species and its habitats. Studies in Avian Biology No. 38. University of California Press. 646pp.

Behney, A. C., C. W. Boal, H. A. Whitlaw, and D. R. Lucia. 2011. Interactions of raptors and Lesser

Prairie-Chickens at leks in the Texas Southern High Plains. Wilson Journal of Ornithology

123:332-338.

Braunisch, V., P. Patthey, and R. Arlettaz. 2011. Spatially explicit modeling of conflict zones between

wildlife and snow sports: prioritizing areas for winter refuges. Ecological Applications 21:955-

967. (Black Grouse).

Budeau, D. 2010. 2010 forest grouse parts collection summary. Unpublished report. Oregon

Department of Fish and Game. 29pp. (Ruffed Grouse, Spruce Grouse, Blue Grouse).

Bunnefeld, N., D. C. Reuman, D, Baines, and E. J. Milner-Gulland. 2011. Impact of unintentional

selective harvesting on the population dynamics of Red Grouse. Journal of Animal Ecology

XXX:XXX-XXX (online early).

Casazza, M. L., P. S. Coates, and C. T. Overton. 2011. Linking habitat selection and brood success in

Greater Sage-Grouse. Pp. 151-168 in B. K. Sandercock, K. Martin, and G. Segelbacher

http://www.suttoncenter.org/pages/publications

http://digitalcommons.usu.edu/nrei/vol16/iss1/5


43

(editors). Ecology, conservation, and management of grouse. Studies In Avian Biology (no.


Caudill, D. 2011. Factors affecting Greater Sage-Grouse (Centrocercus urophasianus) survival and

movement in south-central Utah. M. Sc. Thesis. Utah State University.

Christiansen, T. J., and C. M. Tate. 2011. Chapter eight – Parasites and infectious diseases of Greater

Sage-Grouse. Pp. 113-126 IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-

Grouse – ecology and conservation of a landscape species and its habitats. Studies in Avian

Biology No. 38. University of California Press. 646pp.

Chubbs, T. E., and F. R. Phillips. 2010. Recent range expansion of Ruffed Grouse, Bonasa umbellus, in

Labrador. Canadian Field-Naturalist 124:45–48.

Ciereszko, A., G. J. Dietrich, E. Liszewska, A. Krzywinski, and A. Kobus. 2011. Short-term storage and

cryopreservation of Black Grouse Tetrao tetrix and Capercaillie T. urogallus semen. European

Journal of Wildlife Research 57:383-388.

Connelly, J. W., C. A. Hagen, and M. A. Schroeder. 2011. Chapter three – Characteristics and dynamics

of Greater Sage-Grouse populations. Pp. 53-67 IN: S. T. Knick and J. W. Connelly (Eds). 2011.

Greater Sage-Grouse – ecology and conservation of a landscape species and its habitats. Studies

in Avian Biology No. 38. University of California Press. 646pp.

Connelly, J. W., S. T. Knick, C. E. Braun, W. L. Baker, E. A. Beever, T. Christiansen, K. E. Doherty, E.

O. Garton, E. S. Hanser, D. H. Johnson, M. Leu, R. F. Miller, D. E. Naugle, S. J. Oyler-

McCance, D. A. Pyke, K. P. Reese, M. A. Schroeder, S. J. Stiver, B. L. Walker, and M. J.

Wisdom. 2011. Chapter twenty-four – Conservation of Greater Sage-Grouse – a synthesis of

current trends and future management. Pp. 549-563 IN: S. T. Knick and J. W. Connelly (Eds).

2011. Greater Sage-Grouse – ecology and conservation of a landscape species and its habitats.

Studies in Avian Biology No. 38. University of California Press. 646pp.

Connelly, J. W., E. T. Rinkes, and C. E. Braun. 2011. Chapter four – Characteristics of Greater Sage-

Grouse habitats. Pp. 69-83 IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-



Corman, K. S. 2011. Conservation and landscape genetics of Texas Lesser Prairie-Chicken:population

structure and differentiation, genetic variability, and effective size. M. Sc. Thesis. Texas A &

M University-Kingsville. 166pp.

Davies, K. W., C. S. Boyd, J. L. Beck, J. D. Bates, T. J. Svejcar, and M. A. Gregg. 2011. Saving the

sagebrush sea: An ecosystem conservation plan for big sagebrush plant communities. Biological

Conservation 144:2573-2584. (Greater Sage-Grouse).

Doherty, K. E., J. L. Beck, and D. E. Naugle. 2011. Comparing ecological site descriptions to habitat

characteristics influencing Greater Sage-Grouse nest site occurrence and success. Rangeland

Ecology and Management 64:344-351.

Doherty, K. E., D. E. Naugle, H. E. Copeland, A. Pocewicz, and J. M. Kiesecker. 2011. Chapter twenty-

one – Energy development and conservation tradeoffs – systematic planning for Greater Sage-

Grouse in their eastern range. Pp. 505-516 IN: S. T. Knick and J. W. Connelly (Eds). 2011.



Ehrich, D., J.-A. Henden, R. A. Ims, L. O. Doronina, S. T. Killengren, N. Lecomte, I. G. Pokrovsky, G.

Skogstad, A. A. Sokolov, V. A. Sokolov, and N. G. Yoccoz. 2011. The importance of willow

thickets for ptarmigan and hares in shrub tundra: the more the better? Oecologia XXX:XXX-

XXX (online early) (Willow Ptarmigan).

Erickson, H. J. 2011. Herbaceous and avifauna responses to prescribed fire and grazing timing in a high-

elevation sagebrush ecosystem. Ph. D. Dissertation. Colorado State University. 180pp.

(Greater Sage-Grouse).

Fedy, B. C., and C. L. Aldridge. 2011. The importance of within-year repeated counts and the influence

of scale on long-term monitoring of sage-grouse. Journal of Wildlife Management 75:1022-

1033.

Fedy, B. and K. Martin. 2011. The influence of fine-scale habitat features on regional variation in

population performance of alpine White-Tailed Ptarmigan. Condor 113:306-315.

Freemuth, J. C. 2011. Foreword – Thoughts on the role of science in making public policy. Pp. xiii-xvii

IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and




44

Fritzler, J. M., T. M. Craig, A. Elgayar, C. Plummer, R. S. Wilson, M. J. Peterson, and G. Zhu. 2011. A

new Eimeriid (Apicomplexa) species from endangered Attwater's Prairie Chickens

(Tympanuchus cupido attwateri) in Texas. The Journal of Parasitology 97:671-675.

Garton, E. O., J. W. Connelly, J. S. Horne, C. A. Hagen, A. Moser, and M. A. Schroeder. 2011. Chapter

fifteen – Greater Sage-Grouse population dynamics and probability of persistence. Pp. 293-381




Gavashelishvili, A., and Z. Javakhishvili. 2010. Combining radio-telemetry and random observations to

model the habitat of near threatened Caucasian Grouse Tetrao mlokosiewiczi. Oryx 44:491-500.

Geary, M., A. H. Fielding, and S. J. Marsden. 2011. The anatomy of population change in a Black

Grouse population 1992–2008. Oecologia XXX:XXX-XXX (online early).

Gibson, R. M., V. C. Bleich, C. W. McCarthy, and T. L. Russi. 2011. Hunting lowers population size in

Greater Sage-Grouse. Pp. 307-316 in B. K. Sandercock, K. Martin, and G. Segelbacher



Gregory, A. J. 2011. Landscape genetics and behavioral ecology of Greater Prairie-Chickens

(Tympanuchus cupido). Ph. D. Dissertation. Kansas State University. 142pp.

Gregory, A. J., L. B. McNew, T. J. Prebyl, B. K. Sandercock, and S. M. Wisely. 2011. Hierarchical

modeling of lek habitats of Greater Prairie-Chickens. Pp. 21-32 in B. K. Sandercock, K. Martin,

and G. Segelbacher (editors). Ecology, conservation, and management of grouse. Studies In

Avian Biology (no. 39), University of California Press, Berkeley, CA.

Hagen, C. A. 2010. Impacts of energy development on prairie grouse ecology: a research synthesis.

Transactions of the 75th North American Wildlife and Natural Resources Conference 75:96-103.

(Greater Sage-Grouse, Gunnison Sage-Grouse, Greater Prairie-Chicken, Lesser Prairie-Chicken,

Sharp-tailed Grouse).

Hagen, C. A. 2011. Chapter six – Predation on Greater Sage-Grouse. Pp. 95-100 IN: S. T. Knick and J.

W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and conservation of a landscape

species and its habitats. Studies in Avian Biology No. 38. University of California Press.

646pp.

Hagen, C. A., J. C. Pitman, T. M. Loughin, B. K. Sandercock, R. J. Robel, and R. D. Applegate. 2011.

Impacts of anthropogenic features on habitat use by Lesser Prairie-Chickens. Pp. 63-76 in B. K.

Sandercock, K. Martin, and G. Segelbacher (editors). Ecology, Conservation, and management

of grouse. Studies In Avian Biology (no. 39), University of California Press, Berkeley, CA.

Hamilton, S., and D. Manzer. 2011. Estimating lek occurrence and density for Sharp-tailed Grouse. Pp.

33-50 in B. K. Sandercock, K. Martin, and G. Segelbacher (editors). Ecology, conservation, and

management of grouse. Studies In Avian Biology (no. 39), University of California Press,

Berkeley, CA.

Hancock, M., A. Amphlett, R. Proctor, D. Dugan, J. Willi, P. Harvey, and R, W. Summers. 2011.

Burning and mowing as habitat management for Capercaillie Tetrao urogallus: An experimental

test. Forest Ecology and Management 262:509-521.

Hansen, M. C., C. A. Hagen, T. M. Loughin, D. A. Budeau, V. L. Coggins, and B. S. Reishus. 2011.

Temporal changes in age and sex ratios of forest grouse harvested in northeastern Oregon.

Journal of Wildlife Management XXX:XXX-XXX (online early). (Dusky Grouse, Ruffed

Grouse).

Hanser, S. E., and S. T. Knick. 2011. Chapter nineteen – Greater Sage-Grouse as an umbrella species for

shrubland passerine birds – a multiscale assessment. Pp. 475-487 IN: S. T. Knick and J. W.

Connelly (Eds). 2011. Greater Sage-Grouse – ecology and conservation of a landscape species

and its habitats. Studies in Avian Biology No. 38. University of California Press. 646pp.

Henden, J.-A., R. A. Ims, N. G. Yoccoz, and S. T. Killengreen. 2011. Declining Willow Ptarmigan

populations: The role of habitat structure and community dynamics. Basic and Applied Ecology

12:413-422.

Hess, J. E. 2011. Greater Sage-Grouse (Centrocercus urophasianus) habitat response to mowing and

prescribed burning Wyoming big sagebrush and influence of disturbance factors on lek

persistence in the Bighorn Basin, Wyoming. M. Sc. Thesis. University of Wyoming. 168pp.

Hibbs, W. B. 2011. Assessing wildlife habitat suitability for ecological sites and state and transition

models. Ph. D. Dissertation. Colorado State University. 174pp. (Greater Sage-Grouse).

Hoglund, J. J. K. Larsson, C. Corrales, G. Santafe, D. Baines, and G. Segelbacher. 2011. Genetic

structure among Black Grouse in Britain: implications for designing conservation units. Animal

Conservation 14:400-408.


45

Hull, S., D. Sample, D. Drake, S. Fandel, L. Kardash, O. LeDee, and S. Schwab. 2011. The Wisconsin

Greater Prairie-Chicken program: integrating research, management, and community outreach in

the 21st century. Passenger Pigeon 73:89-99.

Jie, W., F. Yun, S. Klaus, and Y.-H. Sun. 2011. Winter foraging strategy of the Chinese Grouse (Bonasa

sewerzowi): ecological and physiological factors. Journal of Ornithology XXX:XXX-XXX.

(online early).

Johnson, D. H., M. J. Holloran, J. W. Connelly, S. E. Hanser, C. L. Amundson, and S. T. Knick. 2011.

Chapter seventeen – Influences of environmental and anthropogenic features on Greater Sage-

Grouse populations, 1997-2007. Pp. 407-430 IN: S. T. Knick and J. W. Connelly (Eds). 2011.



Kaczor, N. W., K. M. Herman-Brunson, K. C. Jensen, M. A. Rumble, R. W. Klaver, and C. C. Swanson.

2011. Resource selection during brood-rearing by Greater Sage-Grouse. Pp. 169-178 in B. K.

Sandercock, K. Martin, and G. Segelbacher (editors). Ecology, conservation, and management


Kaczor, N. W., K. C. Jensen, R. W. Klaver, M. A. Rumble, K. M. Herman-Brunson, and C. C. Swanson.

2011. Nesting success and resource selection of Greater Sage-Grouse. Pp. 107-118 in B. K.



Kaler, R. S. A., and B. K. Sandercock. 2011. Effects of translocation on the behavior of island

ptarmigan. Pp. 295-306 in B. K. Sandercock, K. Martin, and G. Segelbacher (editors). Ecology,

conservation, and management of grouse. Studies in Avian Biology (no. 39), University of

California Press, Berkeley, CA.

Knick, S. T. 2011. Chapter one – Historical development, principle federal legislation, and current

management of sagebrush habitats. Pp. 13-31 IN: S. T. Knick and J. W. Connelly (Eds). 2011.



Knick, S. T., and J. W. Connelly. 2011. Greater Sage-Grouse – ecology and conservation of a landscape


646pp.

Knick, S. T., and J. W. Connelly. 2011. Greater Sage-Grouse and sagebrush – an introduction to the

landscape. Pp. 1-9 IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-Grouse –

ecology and conservation of a landscape species and its habitats. Studies in Avian Biology No.

38. University of California Press. 646pp.

Knick, S. T., and S. E. Hanser. 2011. Chapter sixteen – Connecting pattern and process in Greater Sage-

Grouse populations and sagebrush landscapes. Pp. 383-405 IN: S. T. Knick and J. W. Connelly

(Eds). 2011. Greater Sage-Grouse – ecology and conservation of a landscape species and its

habitats. Studies in Avian Biology No. 38. University of California Press. 646pp.

Knick, S. T., S. E. Hanser, R. F. Miller, D. A. Pyke, M. J. Wisdom, S. P. Finn, E. T. Rinkes, and C. J.

Henny. 2011. Chapter twelve – Ecological influence and pathways of land use in sagebrush.

Pp. 203-251 IN: S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology

and conservation of a landscape species and its habitats. Studies in Avian Biology No. 38.


Kokott. J. 2011. Schutz der natürlichen Lebensräume und der wildlebenden Arten – Braunbär (Ursus

arctos) – Auerhuhn (Tetrao urogallus). [Protection of natural habitats and of wild species -

brown bear (Ursus arctos) - Capercaillie (Tetrao urogallus).] Natur und Recht 33:666-676. (in

German).

Kukal, C. A. 2010. The over-winter ecology of Lesser Prairie-Chickens (Tympanuchus pallidicinctus) in

the northeast Texas Panhandle. M. Sc. Thesis, Texas Tech University, Lubbock, USA.

Laiolo, P., M. J. Bañuelos, B. Blanco-Fontao, M. García, and G. Gutiérrez. 2011. Mechanisms

underlying the bioindicator notion: spatial association between individual sexual performance

and community diversity. PLoS ONE 6(7):e22724. doi:10.1371/journal.pone.0022724

(Capercaillie).

Lampila, P., E. Ranta, M. Monkkonen, H. Linden, and P. Helle. 2011. Grouse dynamics and harvesting in

Kainuu, northeastern Finland. Oikos 120:1057-1064. (Capercaillie, Black Grouse, Hazel

Grouse, Willow Grouse).

Lebigre, C., R. V. Alatalo, J. Kilpimaa, V. Staszewski, and H. Siitari. 2011. Leucocyte counts variation

and measures of male fitness in the lekking Black Grouse. Journal of Ornithology XXX:XXX-

XXX (online early).


46

Leu, M., and S. E. Hanser. 2011. Chapter thirteen – Influences of the human footprint on sagebrush

landscape patterns – implications for sage-grouse conservation. Pp. 253-271 IN: S. T. Knick and

J. W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and conservation of a landscape


646pp.

Lyons, E. K., R. S. Jones, J. P. Leonard, B. E. Toole, R. A. McCleery, R. R. Lopez, M. J. Peterson, and

N. J. Silvy. 2011. Regional variation in nesting success of Lesser Prairie-Chickens. Pp. 223-

232 in B. K. Sandercock, K. Martin, and G. Segelbacher (editors). Ecology, conservation, and

management of grouse. Studies In Avian Biology (no. 39), University of California Press,

Berkeley, CA.

Mandich, C. A. 2011. Seasonal habitat distribution and parasite survey of Greater Sage-Grouse in

Western Natrona County, Wyoming. M. Sc. Thesis. University of Wyoming. 113pp.

Manley, G. A., M. Lieser, and P. Berthold. 2011. Studies of infrasound production and perception by the

Capercaillie (Tetrao urogallus): a reply to Freeman and Hare. Journal of Ornithology 152:817-

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Martin, K., S. Wilson, and S. J. Hannon. 2011. Mechanisms underlying variation in renesting ability of

White-tailed Ptarmigan. Pp. 233-246 in B. K. Sandercock, K. Martin, and G. Segelbacher



Matthews, T. M., A. J. Tyre, J. S. Taylor, J. J. Lusk, and L. A. Powell. 2011. Habitat selection and brood

survival of Greater Prairie-Chickens. Pp. 179-194 in B. K. Sandercock, K. Martin, and G.

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Biology (no. 39), University of California Press, Berkeley, CA.

McLachlan, M., A. Bartuszevige, and D. Poole. 2011. Evaluating the potential of the Conservation

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McNew, L. B., A. J. Gregory, S. M. Wisely, and B. K. Sandercock. 2011. Reproductive biology of




McNew, L. B., A. J. Gregory, S. M. Wisely, and B. K. Sandercock. 2011. Human-mediated selection on

life-history traits of Greater Prairie-Chickens. Pp. 255-266 in B. K. Sandercock, K. Martin, and

G. Segelbacher (editors). Ecology, conservation, and management of grouse. Studies in Avian

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McNew, L. B., T. J. Prebyl, and B. K. Sandercock. 2011. Effects of rangeland management on the site

occupancy dynamics of prairie-chickens in a protected prairie preserve. Journal of Wildlife

Management XXX:XXX-XXX (online early).

McRoberts, J. T., M. J. Butler, W. B. Ballard, H. A. Whitlaw, D. A. Haukos, and M. C. Wallace. 2011.

Detectability of Lesser Prairie-Chicken leks: A comparison of surveys from aircraft. Journal of

Wildlife Management 75:771-778.

Miller, R. F., S. T. Knick, D. A. Pyke, C. W. Meinke, M. J. Wisdom, and A. L. Hild. 2011. Chapter ten -

Characteristics of sagebrush habitats and limitations to long-term conservation. Pp. 145-184 IN:

S. T. Knick and J. W. Connelly (Eds). 2011. Greater Sage-Grouse – ecology and conservation

of a landscape species and its habitats. Studies in Avian Biology No. 38. University of

California Press. 646pp.

Montadert, M., and P. Leonard. 2011. Biologie de la reproduction de la Gelinotte des Bois Bonasa

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partie). [Breeding biology of Hazel

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Montadert, M., and P. Leonard. 2011. Natal dispersal affects population dynamics of Hazel Grouse in

heterogeneous landscapes. Pp. 89-106 in B. K. Sandercock, K. Martin, and G. Segelbacher



Musil, D. D. 2011. Use of dwarf sagebrush by nesting Greater Sage-Grouse. Pp. 119-136 in B. K.



Naugle, D. E., K. E. Doherty, B. L. Walker, H. E. Copeland, M. J. Hollorand, and J. D. Tack. 2011. Sage-

grouse and cumulative impacts of energy development. Pp. 55-70 in D. E. Naugle (Ed.).

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Naugle, D. E., K. E. Doherty, B. L. Walker, M. J. Holloran, and H. E. Copeland. 2011. Chapter twenty –

Energy development and Greater Sage-Grouse. Pp. 489-503 IN: S. T. Knick and J. W. Connelly



Nelson, A. R. 2010. Ecology of Prince of Wales Spruce Grouse. M. Sc. Thesis. University of Alaska,

Fairbanks. 85pp.

Niemuth, N. D. 2011. Spatially explicit habitat models for prairie grouse. Pp. 3-20 in B. K. Sandercock,

K. Martin, and G. Segelbacher (editors). Ecology, conservation, and management of grouse.

Studies In Avian Biology (no. 39), University of California Press, Berkeley, CA.

Novoa, C., J.-F. Desmet, J.-F. Brenot, B. Muffat-Joly, M. Arvin-Berod, J. Resseguier, and B. Tran. 2011.

Demographic traits of two alpine populations of Rock Ptarmigan. Pp. 267-282 in B. K.



Nudds, R. L., L. P. Folkow, J. J. Lees, P. G. Tickle, K.-A. Stokkan, and J. R. Codd. 2011. Evidence for

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spruce (Picea abies) important in some habitats?] M. Sc. Thesis. Swedish University of

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Oyler-McCance, S. J., and T. W. Quinn. 2011. Chapter five – Molecular insights into the biology of

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Oyler-McCance, S. J., C. A. Strickler, J. St. John, C. E. Braun, G. T. Wann, M. S. O’Donnell and C. L.

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Pp. 283-294 in B. K. Sandercock, K. Martin, and G. Segelbacher (editors). Ecology,

conservation, and management of grouse. Studies In Avian Biology (no. 39), University of


Parker, S., and M. Vasquez. 2011. Flat Top Mountain Gunnison Sage-Grouse habitat monitoring study.

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Patten, M. A., C. L. Pruett, and D. H. Wolfe. 2011. Home range size and movements of Greater Prairie-

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conservation, and management of grouse. Studies In Avian Biology (no. 39), University of


Patten, M. A., and B. D. Smith-Patten. 2011. “As is” philosophy: conservation biology’s real hope.

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Patthey, P., N. Signorelli, L. Rotelli, and R. Arlettaz. 2011. Vegetation structural and compositional

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Capercaillie (Tetrao urogallus): redesigning sexing and microsatellite primers to increase

efficiency on faeces samples. Conservation Genetics Resources 3:483-487.

Pitman, J. 2011. Prairie Chicken lek survey – 2011. Performanc Report, Statewide Wildlife Research

and Surveys, A Contribution of Pittman-Robertson Funds, Federal Aid in Wildlife Restoration,

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Pyke, D. A. 2011. Chapter twenty-three – Restoring and rehabilitating sagebrush habitats. Pp. 531-548





48

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Reese, K. P., and J. W. Connelly. 2011. Chapter seven – Harvest management for Greater Sage Grouse –

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and its habitats. Studies in Avian Biology No. 38. University of California Press. 646pp.

Rojas, M., I. Gonzales, M. A. Pavon, N. Pegels, P. A. Hernandez, T. Garcia, and R. Martin. 2011.

Development of a real-time PCR assay to control the illegal trade of meat from protected

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Ross, J. D. 2011. The evolutionary history, demographic independence and conservation status of two

North American prairie bird species: The Greater Prairie Chicken and the Lark Sparrow. Ph. D.

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Sandercock, B. K., K. Martin, and G. Segelbacher (Eds.). 2011. Ecology, conservation, and management

of grouse. Studies in Avian Biology, No. 39. University of California Press, Berkeley,

California. 376pp.

Sawa, Y., Y. Takeuchi, and H. Nakamura. 2011. Nest site selection and nesting biology of Rock

Ptarmigan Lagopus muta japonicus in Japan. Bird Study 58:200-207.

Schole, A. C., T. W. Matthews, L. A. Powell, J. J. Lusk, and J. S. Taylor. 2011. Chick survival of




Schrag, A., S, Konrad, S. Miller, B. Walker, and S. Forrest. 2011. Climate-change impacts on sagebrush

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Schweiger, A,-K., U. Nopp-Mayr, and M. Zohmann. 2011. Small-scale habitat use of Black Grouse

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49

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Lagopus lagopus scoticus on moors in northern England. Wildlife Biology 17:135-142.

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Environmental Surveillance, Education, and Research Program, Contract No. DE-NE-0000-300.

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captive birds. Systematic and Applied Microbiology XXX:XXX-XXX (online early).

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with English abstract).


50

SNIPPETS

Pyrenean Network for the mountain Pyrenean galliforms -

GALLIPYR ProjectPOCTEFA 2007 -2013 – 01/12/2008 to 30/11/2011

Céline Claustre (Forespir)

Partners:

· FORESPIR – Lead partner

· Govern de ANDORRA

· Office National des Forêts-ONF

· Office National de la Chasse et de la Faune Sauvage-ONCFS

· Fédération Régional des Chasseurs de Midi-Pyrénées-FRC MP

· Generalitat de Catalunya-GENCAT-DMAH

· Centre Tecnológic Forestal de Catalunya-CTFC

· Conselh Generau del Val d’Aran

· Gestión Ambiental Viveros y Repoblaciones de Navarra-GAVRN

· Diputación Foral de Alava

· Observatoire des Galliformes de Montagne - OGM

· Fédérations Départementales des Chasseurs

Co-financing: DREAL Midi-Pyrénées, Conseil Régional Aquitaine, Conseil Régional Midi-Pyrénées,

Conseil Général Pyrénées-Atlantiques, Conseil Général Hautes-Pyrénées, Communautés de Communes

des 3 Vallées, Communautés de Communes du Canton de Saint-Béat, la Généralité de Catalogne, le

Gouvernement de Navarre et la Diputation Foral de Alava, Andorre.

Total Cost: 2 446 940 €

European Funding: 1 534 119 €

Contact: Virginie Fabre - Director of FORESPIR and GALLIPYR Lead Partner - 64 rue Raymond IV -

31000 TOULOUSE – FRANCE Phone: + 33 (0)5 34 41 43 20, [email protected].

Context

The GALLIPYR project is designed to harmonize the methods of monitoring and management of 3

species of mountain galliforms between 3 States that make up the Pyrenean Massif (Spain-France-

Andorra): capercaillie (Tetrao urogallus), rock ptarmigan (Lagopus muta) and mountain grey partridge

(Perdix perdix hispaniensis). Actions are also conducted to encourage the return of hazel grouse (Bonasa

bonasia), extinct species of the Pyrenees following due to human activity pressure. On the French side of

the Pyrenees, methods of monitoring of these populations exist across the Mountain Galliform

Observatory (OGM), the GALLIPYR project will expand and develop expertise between French-Spanish-

Andorran specialists for the mountain game fowl, in the whole of the Pyrenean range for a better cross-

border balance.

The GALLIPYR Project worklines

The project provides for the creation of a network of Pyrenean mountain galliforms, creation of a

database of Pyrenees accessible (with restricted access) from the website of the project about these

species and implementation of actions for the species and their habitats, with, in particular:

actions concerning the three species: realization of inventories of cables and lethal fences and

visualization of a part of it; canalization of human movements on pilot areas of high strategic value for

their preservation to diminish disturbance; monitoring of populations of this species and modeling the

potential habitats for ptarmigan and grey partridge and breeding habitat mapping.

specific worklines for capercaillie: improvement of breeding and wintering habitats in strategic areas

(core areas, corridors), on significant surfaces in regard to the present range; realization of a guide for

forest management adapted at the important diversity of Pyrenean forests, and study to test the

influence of terrestrial predators and the wild boar on Capercaillie populations.



51

specific worklines for rock ptarmigan: a program of translocation of individuals, from a strong

population toward a population genetically weak, with long-term monitoring the participation in the

reproduction of released birds, the evolution of the reproductive success, and an update of the data on

the causes of mortality of Rock Ptarmigan will be also performed.

specific worklines for grey partridge: a practical guide of the mountain and subalpine meadows and

shrublands management modes will be performed, actions to improve the habitat and recovery of

populations in Navarre and the Basque country, where the species disappeared historically, by

translocation of wild birds from strong populations of central Pyrenees.

specific worklines for hazel grouse (Bonasa bonasia): drafting of a plan to reintroduce it, and if the

conditions are met, tests of reintroduction of hazel grouse can be realized in Val of Aran (Spain).

Expected results and perspectives

The actions of visualization of a big number of cables and fences should have a positive impact on the

mortality rate of the mountain galliforms, but also have beneficial effects on a large number of birds

species, including some rare species (owls, raptors,…). For rock ptarmigan: improve the efficiency of the

reproduction and try to reduce a loss of genetic diversity found in a important but marginal population of

this species in the East of the Pyrenees. The methodology of translocation could be transposed to other

species. An increase of the carrying capacity of the habitats of capercaillie and grey partridge is expected

from their improvement, and consequently we hope an positive effect in their demography.

Céline Claustre, Chargée d'études FORESPIR et Animatrice du Projet GALLIPYR, Encargada de

estudios FORESPIR y Animadora del Proyecto GALLIPYR, 64 rue Raymond IV Bât. C, 31000 Toulouse,

[email protected].

Desperately seeking stable, 50-year old landscapes with patches and

long, wide corridors

Conservation corridors are a promising intervention to avoid and mitigate habitat fragmentation and to

allow wildlife to shift their geographic ranges in response to climate change. Conservation scientists and

managers have little evidence that corridors will work because most corridor research has been conducted

at small spatial extents (< 150m corridor length, < 50 m corridor width), in landscapes where the matrix is

a natural landcover (e.g., grassland corridor and patches in a forest matrix), and using response variables

(animal presence or short-term movement) that are weakly related to long-term gene flow and

demographic persistence. To remedy this, we (Paul Beier and Andy Gregory) plan to test the ability of

long (0.5 to 100 km), wide (> 100 m) corridors to maintain gene flow and demographic stability among

populations embedded in human-modified matrix. We need your help to identify suitable landscapes with

corridors and isolated patches. Please visit www.docorridorswork.org for more information on our

project, and to suggest landscapes for our research. We offer a small finder’s fee to those who suggest

sites eventually used in our analysis.

Contact: Dr. Andrew Gregory, [email protected] 1 928 523 2167


http://www.docorridorswork.org/



52

12th

International Grouse Symposium, Matsumoto,

Japan, 20-24 July 2012

IGS News No. 6 (20th

October)

As you know already, the 12th

International Grouse Symposium postponed

1 year because of the March 11 earthquake-tsunami catastrophe and the

nuclear crisis erupted at the Fukushima nuclear power plant in Japan. The

Local Organizing Committee of IGS2012 announces that the 12th

International Grouse Symposium held next year, 20-24 July 2012, in

Matsumoto, Japan.

Seven months has passed after the March 11

catastrophe, Japan has recovered normal way of life

except for the worst-hit coastal regions of Tohoku and

the surrounding areas of the nuclear power plant.

Except just after the earthquake, regulated periodic

reduction in electric power in Tokyo and its

surrounding areas had not done even the peak time of

demand in this summer. We have now no traffic

problems. Japanese government declared Sept. 20 they

will try to achieve a stable condition called “cold

shutdown” of the crisis-hit Fukushima nuclear power

plant by the end of this year. The checks of

radioactive substances in foods are done seriously, so

we have no problems in diets. For these reasons, our

Local Committee decided that it is possible to open

the IGS in the next year.

Almost all our former symposium plans may be kept

and adjustments can be made as needed. The details

will be open in this November in web site.

Hiroshi Nakamura, Faculty of Education, Shinshu University, Nagano, 380-8544, Japan, DESK-PC:

[email protected], Secretary-PC: [email protected], http://cert.shinshu-u.ac.jp/eco_lab/.

Jack Connelly receives Hamerstrom Award for his grouse work

At their fall 2011 meeting, the Prairie Grouse Technical Council presented Jack Connelly with the

Hamerstrom award for his long-term work with prairie grouse. Jack Connelly is working as Principal

Wildlife Research Biologist, Idaho Department of Fish and Game, Northwest Section Representative to

The Wildlife Society, 83 West 215 North, Blackfoot, ID 83221, USA.

Jack has been involved with research and management of sage and sharp-tailed grouse for over

30 years. He is the senior author of the sage-grouse management guidelines and co-author of the habitat

management guidelines for Columbian sharptailed grouse. Jack is co-editor with Steve Knick of the

recently published volume "Greater Sage-Grouse: ecology and conservation of a landscape species and its

habitats." The Hamerstrom Award is named for two of North America's premier early grouse biologists,

Fred and Frances Hamerstrom. The Hamerstroms were a husband and wife team from Wisconsin that

conducted extensive research on prairie chickens and sharp-tailed grouse.



http://cert.shinshu-u.ac.jp/eco_lab/


53

BLM Developing New Sage-Grouse Conservation Strategy

The Bureau of Land Management

(BLM) has announced it will

develop a new range-wide

conservation strategy for the greater

sage-grouse in an attempt to avoid

listing the imperiled species. Recent agreements require the U.S. Fish and Wildlife Service (FWS) to

proceed with listing decisions for 251 Candidate Species, including the Greater Sage-Grouse, over the

next five years. The Service must submit either a proposed listing rule or a “not warranted” finding for the

sage-grouse by 2015.

“This plan is a step in the right direction, but

we still need to know what BLM intends to do for

sage-grouse,” said Steve Holmer, Senior Policy

Advisor at American Bird Conservancy. “What

prescriptions will it include for energy development,

livestock grazing, and cheatgrass control?”

The agencyʻs announcement follows requests

by conservation organizations including ABC,

WildEarth Guardians, Advocates for the West, and

Defenders of Wildlife, as well as FWS and directors of

four western state fish and wildlife agencies that BLM

develop new and improved regulatory mechanisms to

conserve and restore sage-grouse and sagebrush

habitat. FWS found current BLM resource

management plans lacking when it declared the

Greater Sage-Grouse a Candidate Species for listing

under the Endangered Species Act in March 2010.

“To be successful, BLM must heed the

available research that finds that oil and gas drilling,

wind energy development, grazing, roads, and utility

corridors are hazardous to sage-grouse and need better management,” said Dan Casey, ABCʻs Northern

Rockies BCR Coordinator. “Reserve areas for sage-grouse, and other bird species of conservation

concern such as the Sage Thrasher, and Brewerʻs and Sage Sparrows should also be considered in the

plan.”

BLM is now expected to develop multiple environmental impact statements. The public will have

an opportunity to be involved and submit comments on the proposals.

As part of a U.S. Department of Agriculture sage-grouse conservation initiative, $21.8 million

was recently granted to Wyoming farmer and ranches. Funds will be used to tear down fences near

breeding sites, implement sustainable grazing practices, and hire sixteen new staff to help farmers and

ranchers adopt grouse conservation measures.

These measures are urgently needed as a recent study, Factors Associated with Extirpation of

Sage-Grouse by the U.S.D.A. Forest Service and U.S. Geological Survey, found that a variety of human

developments from roads to power lines are having a devastating impact on sage-grouse populations. The

study concludes the birds avoid tall structures such as drilling rigs and communications towers, and found

that abandoned sage-grouse habitat “contained almost 27 times the human density, three times more

agriculture, was 60 percent closer to highways, and had 25 percent higher density of roads.”

Steve Holmer, Senior Policy Advisor, American Bird Conservancy, [email protected].

Greater sage-grouse by Alan Wilson


http://www.abcbirds.org/

http://www.wildearthguardians.org/site/PageServer


54

Gateway West Transmission Line Project a Potential Threat to Sage

Grouse

The Bureau of Land Management (BLM) is in the process of developing plans for the Gateway West

Transmission Line Project, which would create the largest electricity transmission line in the U. S.

According to the draft Environmental Impact Statement (EIS), transmission lines will stretch over 1,100

miles of southern Wyoming and southern Idaho, connecting solar, wind, and geothermal energy

generation sites to population centers. The proposed route coincides with wildlife habitat for sage grouse

(Centrocercus urophasianus), pygmy rabbits (Brachylagus idahoensis), burrowing owls (Athene

cunicularia), and various species of raptor, raising concerns from conservation organizations. Of highest

concern is the imperiled sage grouse, a candidate species for the federal Endangered Species List.

Wyoming has designated 15 million acres in the state as core sage grouse habitat and Idaho has

delineated 9 million acres as key habitat for this species. The Gateway West Transmission Line could

cross through approximately 235 miles of these areas, fragmenting important habitat for the sage grouse.

Local sage grouse working groups are advising the BLM on the issue, and Idaho Power Company and

Rocky Mountain Power have provided the BLM with a plan to mitigate or minimize impacts of the

project on sage grouse, included as an appendix to the draft EIS. The BLM will be holding public

hearings regarding the draft EIS in Wyoming during the first week of October 2011, where the public can

submit comments. Anyone interested in submitting comments on the draft EIS can also do so through a

web form provided by BLM, via e-mail to: [email protected] , or by written mail to:

Bureau of Land Management, Gateway West Project, PO Box 20879, Cheyenne, WY 82003. Comments

are being accepted through 28 October 2011. For more information on the transmission line project,

please visit the BLM project website.

Sources: Bureau of Land Management, E&E Publishing LLC (Landletter)

Reprinted from Wildlife Policy News Volume 21, Issue 10, October 2011 (Editor: Christine

Carmichael [email protected]).

Article on the effect of wind farms on Cantabrian capercaillie.

With the increasing importance of alternative energy sources, wind farms are erected throughout many

parts of Europe. Mountainous areas are particularly attractive for wind parks, but also are the strongholds

for grouse and other threatened species. Thus, conflicts of interest between environmentalists favouring

wind energy and conservationist concerned about threatened species are common. In many places, the

habitats of black grouse, rock ptarmigan, and also capercaillie, are highly suitable for wind parks.

Unfortunately, we lack solid studies on the effects of wind turbines on grouse. From Norway, high

mortality from rock ptarmigan flying into the turbine bases is documented. From central Europe, all we

got are anecdotes. Thus, studies like the one below are filling an important gap, and may play a

significant role in environmental impact studies for wind park development plans.

The Cantabrian capercaillie Tetrao urogallus cantabricus is the most endangered and the only

inhabiting caducifolious forests. Significant differences in number of signs of Cantabrian capercaillie

before and after the construction of the wind farm were found. In a recently described Mediterranean

habitat, the authors assessed capercaillie occurrence before and after a wind farm construction in a

wintering area. After construction of the wind farm signs of capercaillie decreased to zero and space use

changed, likely due to human disturbances derived from the wind farm construction and maintenance.

These changes in habitat use at a local scale and related negative effects on capercaillie are discussed. As

part of the precautionary principle conservation measures for this subspecies should avoid any wind farm

within the current Cantabrian capercaillie range while further research on Capercaillie and wind farms

interactions should be urgently carried out.

González, M. A. & Ena, V. 2011. Cantabrian Capercaillie signs disappeared after a wind farm

construction. - Chioglossa 3: 65-74.

http://www.wy.blm.gov/nepa/cfodocs/gateway_west/draft_eis.html#download

http://www.wy.blm.gov/nepa/cfodocs/gateway_west/documents/deis/Appendix%20C%20-%20Environmental%20Protection%20Plans%20and%20Measures.pdf

http://www.wy.blm.gov/nepa/cfodocs/gateway_west/public_meetings.html

http://www.wy.blm.gov/nepa/cfodocs/gateway_west/public_meetings.html

http://gatewaywesteis.com/?submit-a-comment

http://gatewaywesteis.com/?submit-a-comment


http://www.wy.blm.gov/nepa/cfodocs/gateway_west/

http://www.wy.blm.gov/nepa/cfodocs/gateway_west/

http://www.eenews.net/Landletter/2011/09/08/4



55

New update of The IUCN Red List

The latest update of The IUCN Red List of Threatened Species™ has just been published. It illustrates the

efforts undertaken by IUCN (International Union for Conservation of Nature) and its partners to expand

the number and diversity of species assessed, improving the quality of information in order to obtain a

better picture of the state of biodiversity. With now more than 61,900 species reviewed, another big step

forward has been made toward developing the IUCN Red List into a true ‘Barometer of Life,’ as called

for by leading experts in the magazine Science in 2010. This update offers both good and bad news on the

status of many species around the world. The red list website is http://www.iucnredlist.org/.

The IUCN Red List is not just a register of names and associated threat categories. It is a rich

compendium of information on the threats to the species, their ecological requirements, where they live,

and information on conservation actions that can be used to reduce or prevent extinctions. The Red List is

critical as an indicator of the health of biodiversity, in identifying conservation needs and informing

necessary changes in policy and legislation to drive conservation forward. Red list assessments are

essential for guiding conservation actions.

The text/info was taken from the IUCN press release from 10th

November 2011.

http://www.iucnredlist.org/

grouse news - waconnected.org · there are many interesting articles in this issue of grouse news....

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